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Of the hAT Element Superfamily of Transposable Elements

Integration is peculiar to hAT elements and links them with the process of V(D)J recombination which is responsible, in large part, for the diversity displayed by the vertebrate acquired immune system. This relationship, which is also reflected in the secondary structures of the hAT element transposases and the RAG1 subunit of the enzyme that mediates the recombination of the immunoglobulin and T-cell receptor genes supports the hypothesis that V(D)J recombination and transposable elements, specifically hAT elements, are extant forms of an ancient recombination system, opening up the possibility of using either of these systems as a model for the other. For example, insights into aberrant recombination of V(D)J regions with other regions of the genome might be explored through an examination of hAT element behaviour in more tractable experimental systems.

Transgenic Insects for Biological Control

Genetically transformed insect strains have great potential for improving existing biological control programmes for pest species such as those integrating the SIT, or to develop new control strategies based on the conditional regulation of genes that encode lethal products (Handler 2002a). For beneficial insects, the potential exists to develop transgenic strains having enhanced immune systems, increased longevity and reproductive capacity, or heightened response to odorant cues elicited by prey insects.

Immune avoidance and evasion

May or may not generate a signal (Figure 2 .4). For example, an object or a microorganism with surface binding sites that attach to any membrane-anchored receptors, including GPI-anchored receptors, is internalized by Velcro mechanisms, where the cell membrane is wrapped around the object by adhesive forces (Figure 2 .2). This implies that the recognition of foreign objects and its uptake by cells is determined by the presence of adhesion protein repertoires that are capable of attaching to the surface of the intruding object or organism In this scenario, any objects including those that have not been encountered before by the insect are taken up and internalized as long as receptors bind to it Thus, the interactive mode of uptake can proceed with fortuitous binding activities, while the instructive mode requires specific signals as a prerequisite for cellular uptake reactions . While the two modes of interaction are not mutually exclusive, the interactive process may be a fallback...

Concluding Remarks

Ancestor whiteflies, on the other hand, may have perceived the presence of such large amounts of virus as a potential threat from an alien invader. It is possible that primitive geminivirus-like particles had a deleterious potential far greater than that presented by modern TYLCV-Is. In the absence of an effective immune system, how, if at all, does the insect respond to the deleterious presence of the virus It is possible that the evolutionary trend of the geminivirus-vector partnership was neutralization of any viral function that might negatively impact the insect and its progeny, such as long-term sojourn, replication, invasion of tissues, and transovarial transmission. Mechanisms evolved to ensure the smooth transit of ingested virus through the insect body and its rapid expulsion (transmission to plants). Some features of geminivirus-S. tabaci interaction may be interpreted in the light of these hypothetical evolutionary processes. Two insect strategies might be implemented,...

From immunity to developmental programs a role for pleiotropy

As shown above, PCD has a crucial role in both immunity and development and can be used as an illustrative example of the evolutionary scenarios that may arise from pleiot-ropy Induction of the immune system can have direct or indirect effects on both partners, and most notably on the host developmental program The evolutionary trajectory of the association will depend on the outcome of the modification of the developmental program of the host, which can be (1) negative consequences for the host (2) benefit both for host and symbiont (3) benefit for neither partner (Figure 3 4) Although immune reactions are evoked to protect the host, they can have detrimental effects on various host traits An interesting example is found in Anopheles gambiae where infection by its Plasmodium parasites induces a reproductive cost on host fecundity (Ahmed and Hurd, 2006) In this system, induction of immunity creates an oxidative burst that triggers the apoptotic pathway within the follicular cells,...

Significance Of Fleas

Flea allergy dermatitis (FAD) is a severe condition found primarily in dogs, but also occasionally seen in cats. In a flea-allergic animal, flea salivary antigens initiate a cascade of symptoms, resulting in intense pruritus accompanied by scratching, biting, and self-inflicted trauma. An affected animal typically displays obsessive grooming behavior, with accompanying depilation, leaving the skin with weeping sores, often resulting in secondary infection. FAD is treated with corticosteroids, which possess undesirable side effects, especially when continuous use is required as in chronic FAD cases. Until development of FAD immunotherapy, successful treatment involves flea elimination from the animal's environment and flea bite prevention.

Functions of Hemolymph

Hemolymph is a reservoir of water and chemicals for the body. Hemolymph supplies digested molecules as nutrients and water from ingested water, honey and nectar as well as ions, hormones, and cells of the immune system to muscles and organs surrounding the hemocoel. Hemolymph transports carbon dioxide to be eliminated through the respiratory organs, gut and cuticle. Hemolymph also distributes heat around the body and aids dissipating metabolic heat to the outside. Hemolymph acquires wastes as the breakdown products of metabolism from the surfaces of cells and organs lining the hemocoel. The excretory organs, the Malphigian tubules, remove these wastes from the hemolymph by filtering and passing them into the gut to be excreted with feces. The water of the hemolymph is about twenty percent of the body water of the bee, but percentages vary. In larvae, the hemolymph may hold up to fifty percent of a bee's body water (Chapter 9).

Protection and defense by the hemolymph

Sometimes (c) the actions of predators. In some insects the hemolymph contains malodorous or distasteful chemicals, which are deterrent to predators (Chapter 14). Injury to the integument elicits a wound-healing process that involves hemocytes and plasma coagulation. A hemolymph clot is formed to seal the wound and reduce further hemolymph loss and bacterial entry. If disease organisms or particles enter an insect's body, then immune responses are invoked. These include the cellular defense mechanisms of phagocytosis, encapsulation, and nodule formation mediated by the hemocytes, as well as the actions of humoral factors such as enzymes or other proteins (e.g. lysozymes, prophenoloxidase, lectins, and peptides). The immune system of insects bears little resemblance to the complex immunoglobulin-based vertebrate system. However, insects sublethally infected with bacteria can rapidly develop greatly increased resistance to subsequent infection. Hemocytes are involved in phagocytosing...

The Remaining Endopterygote Orders

This group contains the two largest families of Hymenoptera, ICHNEUMONIDAE (ichneumon flies) (Figure 10.26A) with about 20,000 species (about 3500 in North America), and BRACONIDAE (Figure 10.26B), with some 35,000-40,000 species (2000 in North America). Ichneumonoids are mainly parasitoids oflarvae and pupae of endopterygotes (all orders except Megaloptera and Siphonaptera), laying their eggs either in concealed locations on the host (the primitive condition) or in the host, when various strategies are used to avoid attack by the host's immune system. Venom may be injected into the host prior to oviposition, the poison paralyzing or killing the recipient. Ichneumonids are largely restricted to juvenile stages of endopterygotes, though a few use the eggs of pseudoscorpions and spiders, or adult spiders, as hosts. Many Braconidae are parasitoids of exopterygotes,

The Circulatory System

Recognition of Foreignness and Altered Self. Recognition of an object as foreign or altered self is a key event in phagocytosis, nodule formation and encapsulation, yet how this takes place remains largely a mystery. The limited evidence suggests that there are some similarities to the innate immune system of vertebrates. Thus, for biological foreign objects there appear to be pattern-recognition receptors (PRR) both on the surface of hemocytes and in the plasma. As their name indicates, the PRR recognize, then bind with particular molecular patterns on the surface of the foreign organism. They also recognize altered self, specifically changes to the basal lamina that sits beneath the epidermis (Figure 11.1) and covers all internal organs (Salt, 1970 Lavine and Strand, 2002). This is important at metamorphosis, when juvenile tissues are histolyzed, when an insect is wounded, and when microorganisms penetrate the midgut epithelium. The nature of PRR is poorly known. Several proteins...

Genetics Of Hostparasite Interactions

Intriguingly, other parasitoids use symbiotic association with viruses to attack the immune system of the host. The viruses are either securely integrated in the parasite's genome or end up in the parasite because of feeding in the host. The integrated viruses only replicate in the female parasitoid prior to oviposition. Other parasitoids use virus-like particles (VLPs, e.g. Leptopilina boulardi) to counteract the host immune reaction. VLPs have structures that superficially resemble viruses but that lack nucleic acids (Quicke, 1997). VLPs are used in several ways by the parasitoid. They can form a shield around the parasitoid egg in order to make it invisible for the host immune system (e.g. Venturia canescens). This is a form of molecular mimicry. The VLPs are coded for by the parasitoid genome and for Venturia the gene has been cloned (Theopold et al., 1994). VLPs

Subterranean termite biology and control

Pathogens and their pathogenicity is weak (Culliney and Grace, 2000) . Usually, pathogens are not persistent in the environment of the termite colony because termites employ hygienic measures such as fumigating their nests with naphthalene to kill pathogens (Chen et al , 1998) Termites have the ability to avoid contact with pathogens, remove pathogens through grooming, and isolate infected individuals from the colony (Logan et al., 1990 Culliney and Grace, 2000). Termites also have an efficient immune system to eliminate infections with foreign pathogenic microbes (Rosengaus et al ., 1999) . It has been suggested that even the gut community protects termites against opportunistic invaders and pathogens (Veivers et al , 1982) Overcoming defensive mechanisms and delivering pathogens throughout an entire colony would require a large number of initially infected individuals and a high dose of inoculum, which is difficult to achieve Mass production and reapplications of foreign pathogens...

Virus Symbionin Interactions

Even though the evidence is strong for a role of symbionin in luteovirus persistance, it is important to note that symbionin-luteovirus interactions cannot account for the specificity of luteovirus transmission, because luteoviruses bind with comparable affinity to symbionin from both vector and nonvector aphid species 26, 60 , It appears, rather, that symbionin acts in a nonspecific manner to facilitate the transmission process. The exact mechanism by which this takes place remains unclear. Among the possiblities (which are not mutually exclusive) are (1) symbionin protects the virus from attack by proteases or recognition by the aphid immune system

Evolution of obligate nonnutritional endosymbionts

Genome reduction and genome isolation are linked There is only one possible evolutionary outcome, degradation . Because of genome reduction and isolation, obligate endo-symbionts will have a finite lifespan as a species or a limited shelf life as a functional endosymbiont The provision of the symbiont to the host will eventually degrade to a point were it might severely affect the host In order not to go under with its symbiont, the host must allow for the eventual replacement of the symbiont This means that the host must allow limited concurrent infections with slightly pathogenic bacteria A bacterium has to be infectious to invade a new host species . The bacterium will be naive to the new host and involuntarily cause some pathology The new bacteria have to infect the host intracellularly to escape the immune system

Public Health Importance

On a global scale, the Diptera are the most important order of insects affecting human health Table I), Mosquitoes are the foremost group because of their role as vectors of more pathogenic organisms than any other fly. The adverse impact of malaria, mosquito-borne arboviruses (e.g., yellow fever, dengue, and encephalitis), and metazoan infections such as filariasis on humans worldwide currently exceeds that of recently publicized ailments such as Lyme disease or acquired immune deficiency syndrome. Other human-related vector pathogen relationships involving the Diptera arc exemplified by sand flies and sand-fly fever, bartonellosis, and leishmaniasis by black flies and onchocerciasis (river blindness) and by tsetse and African trypanosomiasis (sleeping sickness), On a global scale, there are about 270 million humans infected with malaria, 90 million with lymphatic filariasis, 17 million with onchocerciasis, and 12 million with

Antimicrobial peptides to kill gut symbionts

Lytic peptides, which are a ubiquitous part of the nonspecific immune system of eukary-otes, disrupt the membranes of bacteria and protozoa by forming channels that lead to cell death (Mutwiri et al , 2000 Boman, 1995, 2003) Because lytic peptides are largely inactive against the electrically neutral, cholesterol-containing cell membranes of higher eukary-otes (Javadpour et al ., 1996 Kamysz et al ., 2003 Boman, 2003), most lytic peptides have low or no toxicity to nontarget organisms such as beneficial insects, humans, and other mammals Because of their mode of action (membrane disruption), the development of resistance mechanisms to lytic peptides is less likely than to chemical insecticides The highly evolved natural lytic peptides, as well as synthetic derivatives, may provide an environmentally friendly alternative to chemical insecticides, because concentrations in the micromolar range are active against microorganisms (Wade et al , 1990) and do not leave toxic residues in the...


Genes involved in host immunity are often also involved in host development (pleiotropy) Hence, any symbiont that is manipulating its host's immune system could potentially also influence its host's developmental program. The intense interactions of the symbiont with the host's immune system, combined with pleiotropy of genes involved in immunity and development, could make host development especially prone to symbiont manipulation This effect is not limited to PCD, but because of its importance for both host immunity and development, PCD might play a crucial role in the evolution of host-symbiont interactions . More generally, pleiotropy between functions could be a major determinant of symbiosis evolution, which can result in several possible evolutionary outcomes However, direct and clear-cut evidence that the mechanisms allowing symbionts to evade the immune system are being used for the modification of the host developmental program is still lacking The main reason for this is...

Review of Hemolymph

Hemolymph receives nutrients from the gut and water and ions from the rectum in the last part of the alimentary canal. Hemolymph supplies water to the organs and tissues, and water consumed in honey and nectar can enter the hemolymph through the alimentary canal. Hemolymph is a reservoir for water for the body. The water of hemolymph holds about twenty percent of the body water of the bee, but its percentage varies. In larvae, hemolymph may contain up to fifty percent of the larva's body water. Hemolymph distributes nutrients, ions and hormones to muscles and organs surrounding the hemocoel and transports cells of the immune system to where they are needed. Numerous hemocytes or blood cells float in the hemolymph. These cells resemble the white blood cells of vertebrates and are of several kinds. Hemolymph distributes digested molecules that are absorbed from the alimentary canal. Hemolymph receives the breakdown products of metabolism that are later removed by the excretory organs,...

Defensive Behavior

Survival and reproduction are the key elements of life. For both elements, defense is a paramount feature without defense, survival, and therefore reproduction, is unlikely. Insects must defend against microorganisms, parasites, and predators and use different strategies against each. The defenses against these attackers differ. The ultimate defense against microorganisms, including bacteria, viruses, protozoans, and fungi, is the immune system. Parasites pose a different challenge. These multicellular organisms live in or on the insect body, sapping vital nutrients and reserves, sometimes damaging essential tissues or organs and causing death. Defenses against parasites are primarily behavioral and life history strategies, with backup from the immune system after parasite attack. Parasitoids are a curious group of attackers that share properties of both parasites and predators. They, like parasites, live in or on the body of the host insect and feed on its blood and tissues. Like...


Hemolymph is the circulating fluid or blood of insects. It moves through the open circulatory system, directly bathing the organs and tissues. Insect hemolymph differs substantially from vertebrate blood, with the absence of erythrocytes and a high concentration of free amino acids being two of the common distinguishing features. The main component of hemolymph is water, which functions as a solvent for a variety of molecules. Water in hemolymph makes up 20 to 50 of the total water in insect bodies, with larval stages generally having a larger relative hemolymph volume than adults. Hemolymph serves as a water storage pool for use by tissues during desiccation and as a storage depot for other types of chemicals. It also contains circulating cells called hemocytes. Hemolymph can function as a hydraulic fluid, for example, in the expansion of a newly molted butterfly's wings. Hemolymph serves important roles in the immune system and in transport of hormones, nutrients, and metabolites.


To avoid the negative selection of foreign pathogens by the termites' hygienic behavior, immune system, and the protective network of the natural gut flora, microbes that are naturally associated with the target insect species (i e , symbionts) could be employed as Trojan horses . According to Greek mythology, the Trojan horse was used to secretly shuttle enemy soldiers into the city of Troy, which was surrounded by impregnable walls Once in the city, the soldiers destroyed Troy and its citizens Modern science explores genetically engineered microorganisms, which serve as Trojan horses to deliver foreign genes into an insect host or an insect population (Beard et al , 2002) The technique of using microorganisms, such as viruses, fungi, or bacterial symbionts, as gene-drive and expression vehicles in a host organism is called paratransgenesis (Durvasula et al , 1997 Beard et al , 1998) An important improvement over foreign biocontrol agents would be that indigenous symbi-onts and...

Unresolved Questions

The role of chemical mediators in the cell-to-cell communication pathways required for successful immune responses as complicated as encapsulation have only just begun to be explored. Also, the initial signaling events that discriminate nonself at the most basic level remain remarkably obscure. This is particularly true for the recognition of eukaryotic parasites, which seem to use a complex mix of strategies such as host immunosuppression, antigen masking, and antigen mimicry to avoid being detected as foreign. Progress in identifying resistance and susceptibility genes in insect hosts, and the complementary virulence avirulence genes in parasites has begun only recently, although efforts are intensifying to clarify these genetic components, particularly in Drosophila. Although insects lack memory cells in their immune system, some long-lived insects such as cockroaches exhibit an enhanced response to a secondary challenge, raising the possibility that insects can show a...


Malathion is much more readily detoxified by humans than by insects, and this differential susceptibility allows the insecticide to be used at rates that kill pests without being proportionately dangerous to mammals. The lethal dose for an adult human would be about 1 xh cups of malathion. Lower doses may suppress the immune system and cause developmental and reproductive abnormalities. By comparison, a closely related compound, para-thion, is deadly enough to have become the weapon of choice for assassins operating within South Africa's apartheid government. After breaking into a residence, a killer would smear the insecticide onto the victim's underwear. The chemical could then enter the body through large hair follicles under the arms and in the crotch Tom Mangold and Jeff Goldberg, Plague Wars The Terrifying Reality of Biological Warfare (New York St. Martin's Griffin, 1999), chap. 23.

Intrinsic Incubation

Mechanisms for tolerating high constant body temperatures and evading the complex immune systems of the vertebrate hosts as well as for tolerating variable body temperatures and avoiding the very different defensive mechanisms of the arthropod vectors. Asexual parasites, such as viruses and bacteria, employ essentially the same life form to infect both vertebrate and arthropod hosts, whereas more highly evolved heterosexual parasites, such as protozoa and helminths, have different life stages in their vertebrate and arthropod hosts. Some asexual parasites, such as the plague bacillus, intermittently may bypass the arthropod host and be transmitted directly from one vertebrate host to another. A disease is the response of the host to infection with the parasite and can occur in either vertebrate or arthropod hosts. Immunity includes all properties of the host that confer resistance to infection and play an important role in determining host suitability and the extent of disease or...