Mini ReviewSalivary glands in ixodid ticks: control and mechanism of secretion
Introduction
Ticks are ectoparasitic, obligate blood-feeding acarines comprised of two major families, the Argasidae and Ixodidae. A third minor family, Nutalliellidae, consists of only one species. Argasid ticks feed rapidly, and consume relatively small meals of blood in contrast to the longer-feeding, more voracious ixodid ticks. The Ixodidae are major vectors of pathogens which cause diseases such as Rocky Mountain spotted fever, Lyme disease, monocytic and granulocytic erhlichiosis in humans and theileriosis and babesiosis in cattle. The prolonged period of attachment and the ability of ixodid ticks to transmit a wide range of pathogens have sparked great interest in studying ixodid salivary gland physiology and salivary gland secretions during tick feeding. Further interest relates to an apparent secretion from a single acinus type (type I) in the salivary glands of free-living ticks that assists in absorbing water from unsaturated air.
Section snippets
Ixodid tick salivary gland secretions
Ixodid ticks are unique amongst ectoparasites in their relatively long-term association with the vertebrate host. Female lone star ticks Amblyomma americanum, for example, attach and feed slowly for 7– 14 days before a 24–48 h period of rapid feeding when they fully engorge and disengage from the host. The female increases from about 4 to 600 mg during this period of attachment. Despite the substantial increase in weight, the quantity of blood imbibed is grossly underestimated if only the
Control of salivary gland fluid secretion
Salivary gland fluid secretion is controlled by nerves, with no evidence of control by hormones (Sauer et al., 1995). Nerves from the tick's central nervous system (synganglion) impinge directly upon the salivary glands (Binnington and Kemp, 1980, Fawcett et al., 1986, Kaufman and Harris, 1983, Megaw, 1977). The nerve(s) run posteriorly along, and are closely associated with, the ducts and terminate in synapses on cells near the lumen of the acini (Fawcett et al., 1986). Three kinds of
Physiology of the salivary glands in unfed ixodid ticks
Ticks often survive months and even years between bloodmeals. A major factor in the tick's ability to survive for prolonged periods without feeding is a remarkable ability to preserve water balance, assisted by an ability to absorb water from unsaturated air (Rudolph and Knulle, 1974, McMullen et al., 1976, Needham et al., 1990, Needham and Teel, 1991, Sigal et al., 1999). The mouth was established as the site of water vapor uptake in ticks (Rudolph and Knulle, 1974) and the salivary glands
Conclusions
The functional diversity of secretory products that facilitate prolonged tick attachment and feeding and the capacity to serve as a conduit for pathogens have prompted detailed studies of the ixodid tick salivary gland. Anatomically, the glands exhibit an extraordinary increase in mass and structural reorganization without an increase in cell number during tick feeding. Plasma membranes and mitochondria in type III and to a lesser extent type II acini proliferate, permitting the salivary glands
Acknowledgements
We thank R.W. Barker and S.K. Wikel for reviewing the manuscript and providing valuable comments. The manuscript was approved for publication by the Oklahoma Agricultural Experiment Station. This work was supported by the Oklahoma Center for the Advancement of Science and Technology (OCAST), grant numbers HR-98-056 and HR-98-057.
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