Pathogens of Invertebrates: Application in Biological Control and Transmission Mechanisms

Biological Control of Parasites
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Includes one chapter on biological control. Attwood, P. BCPC Publ. One of a series of books from the British Crop Protection Council. Covers cultural, chemical and IPM strategies. No refs.

Barbosa, P. Peters, eds. Saunders, Philadelphia, PA. Includes 10, mostly "classic", papers on cultural, biological and integrated methods of insect pest control. Refs after each paper. No index. Schultz, eds. Academic Pr. Beirne, B. Leonard Hill Bks. In his own words: "a collection of opinions and predictions" concerning the future of pest management. Useful background reading for decision makers. Summarizes information on ecology, behavior and life histories of key Canadian crop pests. Refs after each species. Bennett, G. Van Nostrand Reinhold, New York. General, emphasizing household pests.


Invertebrate pathology, like medical and veterinary pathology, for many years has been Application in Biological Control and Transmission Mechanisms. Pathogens of invertebrates: application in biological control and transmission mechanisms by, unknown edition.

Also covers garden and urban insect and vertebrate pest management. Refs after each chapter. Brattsten, L. Ahmad, eds. Plenum Pr. Seven theoretical papers on effects of crop allelochemicals on pest attack and including a useful paper by Raffa on practical applications of this information. Callahan, P. Devin-Adair, Old Greenwich, U. Regarding various structures on insects as "radio transmitters and receivers," Callahan envisages a revolutionary approach to pest control. Carter, W. Wiley, New York. Bioecology of insects as vectors of plant pathogens. Cherrett, J. Ford, I. Herbert, and A. English Univ. Summarizes preventive measures and control of damage by population reduction through use of cultural, inorganic and synthetic chemicals and biological control methods.

Clark, L. Geier, R. Hughes and R. Methuen, Lond. General principles concerning insect numbers and the ecology of pest control from an Australian perspective. Useful discussion of competing theories and systems for classifying control strategies. Davidson, R.

Useful modern summary of the ecology, life habits and control natural, biological, cultural and chemical of key North American pests. Dethier, V. Darwin Pr. A thoughtful discussion of the insect-plant-human relationship with implications for rational action. Drummond, R. George and S. CRC Pr. Emphasis on pesticides. Edwards, C. Useful for general principles; life histories, pictures and keys for British agricultural insect pests. Overemphasizes pesticides. Blackwell, Lond. See especially the review papers on life cycles and population dynamics. A review of the role of diversity in creating stability.

Evans, H. Addison-Wesley Publ. Two chapters provide an excellent summary of key pest control concepts, and IPM. Numerous useful illustrations. Few refs after each chapter. Fenemore, P. Butterworths, Wellington, N. Textbook for students, extension agents, and farmers covering key aspects of pest control using examples from New Zealand.

Frankie, G. Koehler, eds. Refreshing collection of papers on beneficials and pests and ways to control them. Based on a conference in Washington, DC. Fryer, L. Understanding nutrition principles for efficient natural pest control. Gips, T. IASA Publicn. Glass, E. I and II. Office Techn. Covers present status of pest control chemical, cultural, biological, and IPM of wheat, corn, soybean, apple, potato, vegetables and cotton crops; discusses present and future issues; obstacles and actions needed for development and adoption of improved pest control.

I and refs after each paper in Vol. Cambridge Univ. Handbook emphasising pesticides. Describes principles and methods of cultural, biological, and chemical control. Useful summaries of biology, distribution and control of key temperate crop pests. Stresses need to take preventative vs curative approaches. Guilford Pr. Ecological principles population and community dynamics and their application to practical pest management strategies; covers insect-plant interactions, chemical and natural controls biological, plant resistance, genetic, cultural, physical and IPM.

Howard, L. Century, New York. Historical discussion of insects in relation to human activities and how to confront pest problems. Few refs in text. Huffaker, C. Rabb, eds. Includes some useful chapters on applications of ecological knowledge to pest control. Isely, D. PART I. Burgess Publ.

Principles and practical examples of cultural, biological and mechanical methods of pest control. Jones, D. Solomon, eds. Blackwell, Oxford, U. Discusses ecological principles, approaches and application of biological control of weeds, insects, nematodes and vertebrates. Jones, F. Arnold, Lond. A taxonomic review of main British pests of crops and stored grain, pest management, and use and hazards of pesticides.

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Overviews of biologicals, cultural methods, pheromones, plant resistance and implementation of IPM strategies. Kerkut, G. Gilbert, eds. Pergamon Pr. Useful reviews of botanical pesticides and principles of fungal, viral, biological, hormonal and genetic methods of control. Kilgore, W. Doutt, eds. Reviews of biocontrol, microbials, electromagnetism, sterilization, pheromones, repellants, and antifeedants for insects. Kiraly, Z. Szalay-Marzo, eds. Akademiai Kiado, Budapest. Covers defence reactions of plants to infection, insect pest ecology, and new approaches to pest control. Handbook No.

Kuenen, D. Union Conserv. Nature Nat. Brill, Leiden, The Netherlands. Papers on problems associated with pest control. Little, V.

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Optimization of rearing methods for Diaphorina citri Kuwayama, Hemiptera: Liviidae and Tamarixia radiata Waterston, Hymenoptera: Eulophidae , for large-scale production and field establishment of the parasitoid. Green Screen. Nylon monofilament implants were retrieved from the hemocoel and examined using image analysis software to quantify the extent of melanization. As a result, the literature on BC has expanded enormously in both national and international research with the number of publications on insect rearing techniques following the same trend. Morphological abnormalities and cell death in the Asian citrus psyllid Diaphorina citri midgut associated with Candidatus Liberibacter asiaticus. J Infect Dis 4 :

Textbook with pesticide bias. Chapter 19 reviews biological, cultural and chemical controls. Martin, H. Reviews significance of natural chemicals in plant defense against insects. Reviews all types of pest control, but with a strong emphasis on pesticides. Refs throughout.

Matthews, G. Longman, Lond. Emphasizes pesticides, but the visuals are useful for explaining pest control theory. Flint and R. McGraw-Hill, New York. Still the best textbook for information on the biology and life history of key N. American pests. Arranged by commodity and habitat. Useful identification keys based on damage. Overemphasizes chemical control. Refs after each pest provide the only guide to the classic studies of these pests. Luckmann, eds. John Wiley, New York.

Textbook covering ecological and economic aspects of pest management, plant resistance, biological and chemical controls, attractants and repellants. Provides practical examples of pest management programs for cotton, forage, tree fruit, forest and nuisance insects. Miller, J. Miller, eds. Springer-Verlag, New York.

List of diseases spread by invertebrates - Wikipedia

Milne, A. Important review of the theory of natural control. Includes discussion of ecological background underlying pest management, monitoring, and cultural, mechanical, biological and chemical pest control. See also Vol. NAS, Wash. Ethics, economics and principles of pest management; strategies for improvement of chemical controls development of biodegradable pesticides, selective chemicals ; reviews ecological pest control methods: plant resistance, parasites and predators, pathogens, and nonchemical weed control.

Also discusses obstacles for new pest control strategies industry, public agencies. National Academy of Sciences. Studies of pest control problems in corn, cotton, soybeans and timber production. Includes cost-benefit analyses. Refs after each. Reinhold, New York. Primarily on chemical control, but some discussion of resistant plants and insect diseases. Ordish, G. Davies, Lond. A thoughtful history from neolithic times to the present with some future predictions.

Ormerod, E. Useful guide to pre-synthetic pesticide methods of pest control and prevention. Identification of British insect pests. No keys. Refs as footnotes. Pfadt, R. Macmillan Publ. Textbook organized by commodities and emphasising pesticides. Useful for notes on common North American insect pests and for pictorial keys to common families. Numerous plates. Non chemical and environmental controls. Biological pest control of insect pests, plant pathogens, weeds, livestock pests. Dangers of pesticides and pest management systems. Includes many landmark papers by leaders in the USA.

Pimentel, D. Ecological basis for the management of insect populations. Oikos Important conceptual paper. Price, P. Covers trophic relationships, populations, coexistence and competition, communities and distributions; includes a useful discussion on diversity and stability.

Useful background material for ecological approaches to pest control. Pringle, L. Macmillan, New York. Well illustrated elementary introduction to cultural, biological, sterile male and hormonal control methods. Roberts, D. Freeman, San Francisco, CA. Useful reviews of bioecology of crop systems and control of pathogens, nematodes, arthropods, weeds and vertebrates. Scopes, N.

Ledieu, eds. Notes on major British pests and control methods. Strong emphasis on pesticides, but useful concise descriptions of damage characteristics. Singh, S. Ajibola Taylor eds. Worldwide treatment with tropical emphasis, but including useful papers by Irwin and van Emden on N. American control systems for soybean and other grain legumes. Southwood, T. Blackwell Scientific Publ. Background to understanding insect pest outbreaks.

Refs after each article. Stapley, J. Iliffe Bks. Biology of key insect pests from around the world and identification of damage. Overemphasises pesticides. Refs after each pest. Still, W. Iowa State Univ. Basic theory emphasizing plant pathology. Covers cultural, biological and IPM approaches.

Host specificity in biological control: insights from opportunistic pathogens

Tashiro, H. Comstock, Cornell Univ. Summarizes biology, distribution, life history and damage of turfgrass pests: mites, crickets, chinch bugs, some homopteran and other insect pest groups. Also covers turfgrass ecology, vertebrate pests, survey techniques and insect control chemical emphasized. Useful color plates of pests and damage. Teng, P. APS Pr. Paul, MN. Thresh, J. Pitman Publ. Includes 6 papers relating to temperate pests emphasising aphids, carrot fly, and orchard pests.

Trueman, L. Bennett and W. Harvest Publ. Pictorial key, descriptions of biology, damage and control for common household pests. Pesticide bias. Refs after each section. Numerous short papers on pest habits, damage and control. Colour plates of key pests and their damage. Refs after some papers. Varley, G. Gradwell and M. Includes an overview of biocontrol principles.

Wardle, R. Manchester Univ. Covers biological, chemical and cultural controls. Ware, G. Thomson Publ. Includes non-chemical methods for most pests. Watson, T. Moore and G. Concise text covering biological, chemical, physical and cultural controls. Westcott, C. Doubleday, Garden City, NY. Useful descriptions of garden pests and their life histories.

Color pictures of many of them. Overemphasises chemical control. Schoonhoven, eds. June , , Wageningen, Netherlands. Wilson, C. Graham, eds. Academic pr. American pests are introduced. Wilson, M. Broersma and A. Waveland Pr. First of a series; other volumes cover livestock, feed and field crops, vegetable, fruit, ornamentals and household pests; bare essentials of practical insect pest management.

Wood, R. Survey of the harmful effects of modern pest control. Woods, A. McGraw-Hill, Lond. Textbook, semi-technical, modern review of chemical and biological pest control. Rodale Pr. Comprehensive guide to protecting ornamental plants, lawns and vegetables without toxic chemicals. Alford, D.

Wolfe Publ. Control recommendations tend to be chemical. Brook, A. Mitchell Beazley Publ. Useful pictures for identification of pests and diseases of fruits, vegetables and ornamentals. Cravens, R. Non technical, general information about controls non toxic, cultural and chemical. Excellent color photos of insect pests and their controls. Farbenfabriken Bayer Ag. Chemagro Corp. Kansas City, MO. Excellent colour pictures of major pests and damage from around the world. II includes index and lists chemicals.

Harris, K. Collins, Lond. Very useful guide to British pests, many of which occur in N. Johnson, W. An illutrated practical guide. Life histories and color plates of insects and their damage on conifers, evergreens and deciduous plants. Kono, T. State of Calif. Mother Earth News Garden Special 1 2 Useful color photos of harmful and beneficial insects at different stages in their life cycle, with non-chemical methods of control; also covers harmful and helpful vertebrates, such as cats, moles, skunks and bats.

Smith, M. Non-technical; clear, useful colour photos for quick identification of adults and immatures of key N. American insect pests. Golden Pr. Color pictures for quick identification of common N. American insect pests of livestock, gardens, ornamentals, field crops, vegetables and stored products. Apple, J. Smith, eds.

Insect Vectors and Plant Pathogens

Covers practical, economical and philosophical aspects. Useful examples of IPM in different farm systems. Burn, A. Coaker, and P. Jepson, eds. Useful reviews of monitoring, cultural, biological and chemical control methods for pests of key crops, greenhouses, forests and stored products. Conway G. State-of-the-art modeling methods with case studies from agriculture, forestry and health.

Discusses policy options. Extensive use of mathematics. FAO, Rome. Discusses the need for IPM and reviews past efforts throughout the world. Covers history and economics of pest control; practical techniques , case histories and future of IPM. We therefore compared several of these parameters in G. Larvae of a melanic phenotype of the Greater wax moth, Galleria mellonella, were selected for resistance to B. Survival assays conducted on cohorts of the selected S and non-selected NS lines revealed significantly increased resistance survival times of the 25 th generation of the S line to B.

Importantly, this S line did not show statistically significant cross-resistance to M. The defense responses of the S and NS lines to B. Mortality rate of selected line and non-selected line of G. However, during the early stages of topically-applied fungal infection, the cuticular PO activity in infected S, but not infected NS larvae, became elevated above uninfected larval levels at 24 h post inoculation pi for both B. This coincides with peak fungal germination and penetration activities. Cuticular phenoloxidase PO activity A , hemolymph phenoloxidase B , lysozyme-like C activity and encapsulation responses D in hemolymph of G.

Lysozyme-like activity was unchanged in all samples, irrespective of the insect line or treatment Figure 2C. However, this activity was significantly lower 24 h pi with B. Unlike B. Expression of seventeen immunity and putative stress management genes was investigated in the integument and fat body of uninfected control and fungal infected insects from both the S and NS lines. The expression data were complex, with some genes behaving differently under each experimental parameter.

However, several important trends can be reported. The majority of the studied genes were expressed at a lower basal level in uninfected control S larvae compared with the NS larvae, in both the integument and fat body. However, a group of genes coding for four putative stress-management factors Contigs , , , and one AMP Galiomicin exhibited different expression patterns, being slightly higher expressed in the integument but not the fat body of control S larvae compared with control NS insects 1.

Basal expression in uninfected S larvae A is illustrated as a fold change relative to NS uninfected larvae and the x-axis represents basal expression in NS larvae i. Fold induction in infected S larvae is calculated relative to the S uninfected expression B, C. Following topical fungal infection, the pattern of gene expression in the S-line insects infected with B. In the other groups i. NS insects infected with either fungal species, and in S insects infected with M.

The expression of most stress-management and immunity-related genes in the integument of these infected insects was also either unchanged or downregulated, although some AMP genes were upregulated in the integument of M. In contrast, in S-line insects infected with B. Furthermore, differences between gene expression in the two tissue locations was significant, i. Upregulation of AMP genes Gallerimycin, Galiomicin, Gloverin, Cecropin-D, 6-tox and Transferrin a siderophore was 5—times higher in fat body of NS and S fungi infected larvae compared with expression in integuments of infected insects Figure.

Selected insects with enhanced resistance to B. Both NS and S lines exhibited reduced fecundity as a consequence of melanism but this was at equivalent levels in both lines summarized in Table S2. The present study shows that insects can develop resistance to insect pathogenic fungi but the resistance is not absolute and is at a cost. A melanic morph of G. A combination of mechanisms was identified that can account for this resistance, some of which were specific for B. Both the S resistant and NS susceptible lines of this morph expressed a wide repertoire of inducible immune and putative stress management genes.

In the S line these resources were re-focussed to the integument.

Disease prevention, control, and eradication

By concentrating its energies to the first and most important barrier to infection, the host delayed pathogenesis by B. Similarly, infected S line insects exhibited higher cuticular PO activity but not plasma PO or lysozyme. It is possible the S line had evolved an enhanced Beauveria recognition apparatus allowing for a more coordinated and targeted response which would account for the increased activity in the integument but subdued activity in the fat body, the main hemopoietic organ.

The fat body occupies a large portion of the insect hemocoel and is the principle site for the synthesis of AMPs in insects exposed to immunogens, including pathogens, irrespective of point of contact or entry [32] , [33] , [34]. As far as we are aware, there are to date no reports of pathogen specific defense mechanisms involving re-allocation of systemic resources to localised tissues under or at risk of attack. This study shows elevated integumental expression of all but two of the examined putative stress management genes i. The fact that so many putative stress related genes were upregulated signifies their importance in defense responses yet their role is poorly understood and often overlooked.

By working in concert with the AMPs they probably mitigate damage and initiate repair resulting in the increased resistance observed in the S line to B. The importance of the putative stress-response genes is further emphasized in the M. The current study suggests that the resistance in the S line is heritable and multi-factorial, comprising several different physiological traits prioritised not only in terms of location but also timing as they were activated concomitant with the period of fungal penetration of the integument. For example, the elevated cuticular PO activity would inhibit fungal growth through synthesis of melanin and its precursors and through melanin partially shielding cuticular proteins from degradation by Pr1, a major cuticle degrading protease and virulence determinant produced by B.

Genes coding for antioxidants are also highly represented under the same conditions and may defend the host against reactive oxygen species generated by PO activity during cuticle penetration [41]. Interestingly, cecropins do have antifungal activity but have been reported as ineffective against B. Current knowledge is very limited regarding the role in G.

Biocontrol of Invertebrate Pests Using Natural Agents

Gloverin is not currently known to have any potent antifungal activity, however, putative antifungal and antimicrobial peptides may fulfil different and currently unrecognized roles in different hosts and under specific infection conditions. In isolation, each of these responses are unlikely to sufficiently impart the observed resistance, however, their impact may be amplified by a synergy between these and other as yet unidentified traits.

There is growing evidence that insects can acquire long-term protection against pathogens through immune priming or transfer from the parent to the offspring, a phenomenon referred to as transgenerational immune priming [43] , [44]. The current study shows the 25 th generation S line larvae exhibit specific, enhanced resistance to B. An earlier but weaker resistance was also observed in the S line which suggests a heritable and amplified immunocompetence. These observations warrant further investigations of possible underlying genetic and epigenetic mechanisms of resistance.

Melanism is strongly correlated with general pathogen resistance with this trait often being accompanied by a trade off in fecundity, development time and even expression of selected immune components such as lysozyme [22] , [23] , [29] , [46]. In the current study, the pupal weight of the melanic S and NS lines which is directly correlated with fecundity was similar and already pushed close to minimum.

It could be argued that resistance in S line insects would increase further with time but to meet the increased demand on resources may result in untenable sacrifices. This is in marked contrast with insects developing resistance to synthetic chemical insecticides where a slight change can have a profound effect [47] , [48] , [49] whereas resistance to B.

Increased insect resistance to a strain of B. Firstly, we show that there is no cross resistance to other fungi so the extra investment in defense offers no benefit against other pathogens introduced or natural. Secondly, the investment in defense is at the expense of fecundity. Thirdly, the downregulation of the AMPs will probably predispose the insect to opportunistic microbial pathogens. In conclusion, this work reports a previously overlooked adaptation strategy of an insect to a widespread, natural pathogenic fungus. This directional selection with B.

However, the less fecund insects are probably at no evolutionary advantage in the wild, and we postulate that the risk is small of fungal biological control agents failing in the field. The starting population was separated into two lines: one line was exposed to the insect pathogenic fungus Beauveria bassiana and selected for increased resistance to the pathogen S line , and the other line was the non-selected untreated control NS line. The pertinent phenotypic attributes of these lines are summarized in Table S2. The defense responses of the 25 th generation of the S and NS insects to B.

Full details of insect rearing and selection are provided in the Text S1. The susceptibility of S and NS lines to B. Unlike the previous generations, insects from the 25 th generation were not exposed to fungi until they were used for these experiments. Each insect was dipped in an aqueous suspension of the pathogen for 10 s using a concentration of 7.

To determine if there was cross-resistance to other species of fungal pathogen, repeat assays were performed as above, using the insect pathogenic fungus Metarhizium anisopliae isolate P Larvae were observed daily for 10 days up to pupation for both B. The emergent adults were monitored for several days to see what percentage were infected with the pathogen.

All dead insects were removed and examined to confirm the cause of death. All insects used in these experiments were 6 th instar larvae raised in the same cohort. The experiment was repeated independently three times. The total number of individuals used from each line was for the B. Uninfected insects were used as a control. Full details are provided in the Text S1. To determine melanotic encapsulation responses to fungal infection, encapsulation assays were performed in NS and S larvae.

Nylon monofilament implants were retrieved from the hemocoel and examined using image analysis software to quantify the extent of melanization. Full details are provided in the Text S1 and experiments were repeated independently three times. Antibacterial activity in hemolymph plasma was determined by a zone-of-clearance assay using freeze-dried Mirococcus lysodeikticus as a substrate suspended in agarose.

The expression of a range of G. Seventeen target genes were investigated, coding for the antimicrobial peptides gallerimycin, galiomicin, gloverin, cecropin D and 6-tox, the siderophore transferrin, the insect metalloproteinase inhibitor IMPI , one linked to immune signaling Contig , three coding for heat-shock proteins HSP, contig and whose activities ameliorate stress [51] , two coding for enzymes dealing with oxidative stress Contigs and , one linked to G-protein coupled receptor activity and stress response Contigs , one involved in anti-apoptosis activity Contig and two involved with cell proliferation Contigs and Full details are provided in the Text S1 and Table S3.

The following life history traits were monitored in NS and S insects: larval development time from egg hatching to onset of pupation and pupal weight. Data analyses were performed using GraphPad Prism v4. Data were checked for normal Gaussian distribution using the Agostino-Pearson omnibus test, and if abnormally distributed a more conservative non-parametric analysis was applied. Larvae from bioassay experiment uninfected with the fungi all survived the duration of the experiment, for this reason, the uninfected treatment was ignored in the statistical analysis.

Differences in life history traits were compared by non-parametric t-test Mann-Whitney. AMP gene expression in integuments of infected insects. Expression of antimicrobial peptide genes and other immunity genes in the integument of non-selected NS and selected line S larvae after topical B. Expression of genes was assayed in integument tissue by Q-PCR in uninfected insects, and in insects at 24 h after topical infection.

Basal expression in uninfected S larvae bar 1 is calculated as a fold change relative to NS uninfected larvae. Stress-management gene expression in integuments of infected insects. Expression of putative stress-management genes in the integument of non-selected NS and selected line S larvae after topical B. Gene expression was assayed in integument tissue by Q-PCR in uninfected animals, and in animals at 24 h after topical infection. AMP gene expression in fat body of infected insects.

Expression of antimicrobial peptide genes in fat body of non-selected NS and selected line S larvae after topical B. Expression of genes was assayed in fat body tissue by Q-PCR in uninfected animals, and in animals at 24 h after topical infections. Stress-management gene expression in fat body of insects. Expression of putative stress-management genes in fat body of non-selected NS and selected line S larvae after topical B.

Susceptibility of G. Susceptibility of Galleria mellonella larvae of selected and non-selected lines to topical fungal infection with Beauveria bassiana 7. Attributes of melanic and non-melanic G. Attributes of selected resistant and non-selected susceptible melanic morphs of 5th instar Galleria mellonella larvae compared with a non-melanic morph. Mega-analysis of Q-PCR data. Summary showing trends in gene expression in S and NS line G. The authors gratefully acknowledge the helpful advice of Dr. Dan Forman and Prof. Norman Ratcliffe of Swansea University, and Dr.

Markus Rantala of Turku University. Browse Subject Areas? Click through the PLOS taxonomy to find articles in your field. Abstract Microevolutionary adaptations and mechanisms of fungal pathogen resistance were explored in a melanic population of the Greater wax moth, Galleria mellonella. Introduction Insects are predominantly dependent upon cuticular, humoral and cellular defenses to resist fungal pathogens. Results Selection and Insect Survival following B. Download: PPT.

Figure 1. Susceptibility of insects selected by B. Figure 2.