The Circulatory System

The innate immune system of insects is divided into humoral and cellular defense responses. Humoral defenses include antimicrobial peptides, the cascades that regulate coagulation and melanization of hemolymph, and the production of reactive intermediates of oxygen and nitrogen. Cellular defenses refer to hemocyte-mediated responses like phagocytosis and encapsulation. In this review, we discuss the cellular immune responses of insects with emphasis on studies in Lepidoptera and Diptera. Insect hemocytes originate from mesodermally derived stem cells that differentiate into specific lineages identified by morphology, function, and molecular markers. In Lepidoptera, most cellular defense responses involve granular cells and plasmatocytes, whereas in Drosophila they involve primarily plasmatocytes and lamellocytes. Insect hemocytes recognize a variety of foreign targets as well as alterations to self. Both humoral and cell surface receptors are involved in these recognition events. Once a target is recognized as foreign, hemocyte-mediated defense responses are regulated by signaling factors and effector molecules that control cell adhesion and cytotoxicity. Several lines of evidence indicate that humoral and cellular defense responses are well-coordinated with one another. Cross-talk between the immune and nervous system may also play a role in regulating inflammation-like responses in insects during infection. Show more

The insect immune system serves as a key defense against attack by parasitoids. Incompatible hosts often eliminate parasitoids by encapsulation, a process in which hemocytes form a multilayered envelope around the invading organism. Capsule formation involves cooperation between one or more classes of hemocytes and is likely mediated by cytokines and adhesion molecules. Reciprocally, parasitoids have evolved a variety of strategies for overcoming host immune responses. Some parasitoids passively avoid elimination by developing in locations inaccessible to host hemocytes or by possessing surface features that fail to elicit an immune response. Other species actively disrupt the host immune system by injecting specific factors into the host at oviposition. In particular, polydnaviruses associated with several taxa of parasitoids disrupt capsule formation by killing hemocytes or altering their ability to adhere to foreign surfaces. These symbionts have likely played a critical role in evolution of host range and in defining parasitoid-host compatibility. Show more