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Chemical Ecology: The Chemistry of Biotic Interaction
manifestations of parasite-induced or -directed host manipulation are quite complex and vary significantly according to the specific parasitic wasp and host insect species; these changes are referred to as "immunosuppression" and "developmental arrest" (11, 12).
IMMUNOSUPPRESSION AND DEVELOPMENTAL ARREST
The endoparasite's strategy for survival involves avoidance of host recognition as foreign or inhibition of the ensuing immune reaction (12-17). Since encapsulation is apparently the major host defense against parasite egg invasion, host immunosuppression is usually attributed to the wasp's ability to avoid or suppress encapsulation. For the endoparasite Campoletis sonorensis Edson et al. (18) showed that purified viable polydnavirus was responsible for suppressing the host insect's (Heliothis virescens) ability to encapsulate the wasp egg. Theilmann and Summers (19) also showed that both immunosuppression and developmental arrest are induced in H. virescens by purified polydnavirus. Inhibition of encapsulation also correlated with a decrease in circulating hemocytes and reduced plasmatocyte attachment and "spreading" (20).
Upon recognizing a foreign object, hemocytes form a cellular sheath (capsule) around that object and neutralize it (2). Although the identity of all hemocyte classes involved in the encapsulation process is not clear, most of the attention in investigating polydnavirus-induced immunosuppression has been given to the role of granulocytes and plasmatocytes (2). Granulocytes and plasmatocytes are the two principal hemocyte classes known to participate in forming the capsule (14). Granulocytes are thought to be the first hemocyte cell type to reach the foreign object (13, 14). Upon recognizing the invading object—for example, an endoparasite egg—granulocytes degranulate, releasing chemotactic factors that attract more granulocytes and plasmatocytes. Plasmatocytes attach and spread on the surface of the foreign body, for example an endoparasite egg. The role of the host prophenol oxidase cascade and/or other humoral factors in recognition of the endoparasite egg and initiation of encapsulation is not well defined (13).
DIVERSE STRATEGIES FOR SUCCESSFUL PARASITIZATION
In the general spectrum of host-parasite relationships, there is considerable parasite-induced host variability resulting in perturbed host physiology, biochemistry, and developmental behavior (8, 21, 22). There is a wide range of speculation that the parasite has the capacity to modulate or regulate host systems (8, 10, 23). It is clear that the C. sonorensis polydnavirus induces developmental arrest in H. virescens larva (18, 24,