Functional morphology of secretion by the large wax glands (Sensilla Sagittiformia) involved in tick defense

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Abstract

Ticks are protected against ants by release of an allomonal defense secretion from the large wax glands (or type 2 glands) that line their bodies. To explore how the large wax glands operate, before and after microscopic observations of these glands (nonsecreted versus secreted test groups), mass determinations were made for Rhipicephalus sanguineus that had been exhausted of secretion by repeated leg pinching to simulate attack by a predator. Prior to secretion, the glandular organ is fully intact histologically and matches the sensillum sagittiforme, a key taxonomic structure described in the 1940s. The large wax gland is innervated and responds to pressure stimulation as a proprioceptor that stimulates the secretory response. Histological observations after secretion has occurred show that the entire glandular contents and associated cells are jettisoned out of the gland like a syringe. The glandular cellular components are subsequently rebuilt by underlying hypodermal cells within a few days so that secretion can take place again. Presumably, the active allomonal ingredients (hydrocarbons) are released when these derived epidermal cells reach and burst onto the cuticular surface. Our conclusion is that the large wax glands are holocrine and feature intermittent regeneration. Copyright © 2009 Jay A. Yoder et al.

Figures

  • Figure 1: Representative drawings of the large wax glands (sensilla sagittiformia = arrow-organs) in nonfed adult females of R. sanguineus. Top: direct view looking down on the pore opening of the gland. Bar = 10µm. This transverse view of the gland opening shows the fringe (Fr), the spongy elastic chitinous frame (F), the border of the pore opening (B), and the top of the valvular arm (V), that make up the roof of the tuft chamber. Bottom left: redrawing of the original 1942 description of the sensillum sagittiforme in sagittal view from Hyalomma marginatum Koch by Schulze [2] (also redrawn in Dinnik and Zumpt [1]). Bottom right: drawing of the large wax gland from the present study (redrawn from Figure 2(a)). Labeled structures: pore opening of gland (Po), distal arrow-shaped duct (D), valvular arm of the roof of the tuft chamber (V), tuft (T), tuft chamber (Tc), anchoring filament of the tuft (Fa), knot of the scolopale (K), spongy elastic chitin (S), projecting edges of the scolopale (P), proximal terminal chamber (Pc), axial sensory nerve fiber (A), hypodermis (H), glandular cells (G), and enlarged juxtapositioned transdifferentiating epithelial cells (∗). Bar = 30µm.
  • Figure 2: Micrographs of sagittal sections of large wax glands (sensilla sagittiformia = arrow-organs) in nonfed adult females of R. sanguineus. (a) Intact large wax gland showing regular microscopic anatomy (corresponding drawing in Figure 1 with parts labeled that match Schulze’s [2] description). The gland has a restricted pore opening (inset, direct view), a thick spongy chitinous layer, and three large glandular cells occupying most of the proximal terminal chamber lumen. Bar = 30 µm, and Bar = 10 µm (inset). (b) Evacuated large wax gland of ticks that had been stimulated to secrete. Most of the morphology of the large wax gland has been destroyed. Glandular cells have been forced up into the tuft chamber and distal duct, and as they existed under high pressure these cells have been broken into pieces (long arrow). The glandular cell pieces have forced open the pore and widened its diameter and the spongy elastic chitinous frame around the gland pore (inset). Note that juxtapositioned cells are hypertrophied and have started the differentiation process to replace glandular cells that had been ejected during release of the secretion (short arrow). Similar morphology was observed in 20 replicates, and these two micrographs are representative of those replicates. Bar = 30 µm, and Bar = 10 µm (inset).
  • Table 1: Number of days required for the large wax glands to replenish secretion by different nonfed stages of R. sanguineus (93% RH, 25◦C) based on restimulation (leg pinching) following initial exhaustion of secretory reserves. L, larva; N, nymph; AM, adult male; AF adult female; dead tick, killed by freezing and then thawed to room temperature (shown); ∗, results from HCN-killed were similar. Data are mean + SE and values followed by the same superscript letter within a column are not significantly different (ANOVA; P < .05). (N = 3 replicates of 10 ticks for each time point with each replicate coming from a separate rearing batch of ticks).

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CITATION STYLE

APA

Yoder, J. A., Benoit, J. B., Bundy, M. R., Hedges, B. Z., & Gribbins, K. M. (2009). Functional morphology of secretion by the large wax glands (Sensilla Sagittiformia) involved in tick defense. Psyche (London). https://doi.org/10.1155/2009/631030

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