Functional morphology of Gené's organ inHaemaphysalis longicornis (Acari: Ixodidae)

The structure and function of Gené's organ, which secretes the egg wax, were investigated inHaemaphysalis longicornis. The hatching rates of eggs deposited before and after the prevention of the eversion of Gené's organ were compared at the conditions of 30°C, 100% RH and soaking in liquid paraffin. The movement of Gené's organ was observed, and the structures of this organ in unfed and ovipositing females were examined with both light and scanning electron microscopes.The hatching rates of eggs without contact with Gené's organ were markedly lower (4.9%) than that of normal eggs (94.5%) even at the suitable condition for eggs (30°C, 100% RH). In contrast, the hatching rates of eggs soaked in liquid paraffin without contact of Gené's organ were considerably higher (71.8%) than that of eggs without both Gené's organ contact and soaking in liquid paraffin.When oviposition commenced, the camerostome of the female was widened by a downswing of the capitulum, the paired horns of Gené's organ emerged through it, and the oviposited egg from the genital aperture was coated with egg wax. At the same time of retraction of the horns which grip an egg, the capitulum was swung up and the egg was loaded on the scutum.Gené's organ included the outer and inner cuticles, and the outer one is highly folded like bellows when the horns of this organ were retracted into the haemocoel. The undeveloped glands lying in the peripheral region of Gené's organ in the unfed stage commenced their enlargement at the time of the onset of feeding, and they became large tubular glands in the 1-day ovipositing female. The columnar glandular cells of the tubular glands extended their domed apical region into the lumen and contained numerous secretory granules during oviposition.These results suggest that the tubular glands secrete the egg wax which is indispensable for egg survival by serving as a waterproofing agent.     

The effects of precocene treatment on egg wax production in Gené's organ and egg viability in the cattle tickBoophilus microplus (Acarina ixodidae): An ultrastructural study

Was synthesis by Gené's organ, the egg waxing organ of ticks, is disrupted by precocene treatment of engorged femaleBoophilus microplus, resulting in desiccated nonviable eggs due to the absence of a waterproofing wax layer. Electron microscopy shows that precocene has a destructive effect on the glandular cells of Gené's organ. The precocene also inhibited in vitro was synthesis by the gland cells, indicating a selective cytotoxic effect. Oogenesis and oviposition were otherwise unaffected, also suggesting that precocene affects the gland cells directly, rather than indirectly by antagonism of juvenile hormone.     

Morphology and structural organization of gené's organ in Dermacentor reticulatus (Acari: Ixodidae)

Scanning and transmission electron microscopical investigations revealed that Gené's organ in unfed and ovipositing females of Dermacentor reticulatus is formed as a double-sac-structure consisting of an outer epithelial and an inner cuticular sac. In ovipositing ticks the latter emerges through the camerostomal aperture to the exterior. Gené's organ in unfed ticks consists of a corpus, two posterior horns and a pair of undeveloped glands at each side, which differentiate in ovipositing ticks to compound, branched tubular glands with a main efferent duct for each gland opening into the lumen between the epithelial and the cuticular sac. Gené's organ of egg-laying females corresponds basically in morphology and structural organization to that of unfed ticks. Compared with unfed ticks, however, in ovipositing ticks the corpus and horns are longer and broader, the glands are fully developed and the cuticular sac is evertable. The epithelial sac as the outermost part of Gené's organ is continuous with the hypodermis of the basis capituli and the scutum, arises at the camerostomal aperture, forms the corpus and the two blind-ending horns, passes into the epithelium of the main excretory ducts of the glands and envelops the cuticular sac. The cuticular sac passes into the cuticle of the basis capituli and the scutum, arises at the camerostomal aperture, is folded, expands into the horn tips and consists inwards of a smooth epicuticula and outwards of a fibrous endocuticula. Muscles originating from the scutum pass caudomedially through the epithelial sac and are inserted into the cuticular sac. The entire surface of the maximally everted cuticular sac is covered with an amorphous mass. In cleaned samples, ledge-like structures appear on the lateral surface. These ledges turn into balloon-like structures which extend over the medial and dorsal surface. The entire surface including the balloon-like structures and the ledges are provided with numerous cribrate pits.     

An ultrastructural and physiological investigation of the retractor muscles of Gené's organ in the cattle ticksBoophilus microplus andAmblyomma variegatum

The ultrastructures of the retractor muscles of Gené's organs in the cattle ticksBoophilus microplus andAmblyomma variegatum are described. The innervation, neuromuscular junctions, and insertions of the muscles are also described. The retractor muscles are important in controlling the actions of Gené's organ, the egg waxing organ in ticks, during oviposition in the female. The ultrastructural features of the muscles are typical of arthropod muscles, and the nerve terminals at the neuromuscular junctions contain small electron lucent synaptic vesicles with a diameter of 50 nm, and also larger dense core vesicles with a diameter of 100 nm. Evidence is presented implicatingl-glutamate andl-aspartate as putative excitatory transmitters at the tick neuromuscular junction. The excitatory post-synaptic potentials recorded in the muscles were abolished in the presence of low concentrations ofl-glutamic acid andl-aspartic acid, but were unaffected by acetyl-choline, 4-aminobutyric acid and octopamine, suggesting that glutamic acid and aspartic acid interact with receptors on the muscle membrane.     

The effect of avermectins on feeding,salivary fluid secretion,and fecundity in some ixodid ticks

We tested the effects of the potent acaricides, avermectin B₁a (AVM) and 22,23-dihydroavermectin B₁ (ivermectin; IVM) when injected directly into partially fed and fully engorged female ticks. When injected into small ticks (Amblyomma hebraeum Koch), neither drug (up to 100 g/kg b.w.) inhibited subsequent engorgement nor affected oviposition latency, weight of total egg mass laid nor viability of laid eggs. At higher concentrations (1000 and 5000 g/kg b.w.), AVM and IVM were markedly toxic. When injected into engorged ticks, both drugs increased oviposition latency, and reduced fecundity at about 75–100 g/kg b.w. Vitellogenesis, as assessed by a spectrophotometric assay of the ovaries, was not inhibited. Also at 50–100 g/kg b.w., AVM and IVM caused paralysis of the abdominal dorso-ventral muscles and the leg muscles. Both drugs, at 7 days post-injection, proved detrimental to salivary gland function in both small and large ticks, but had little effect on salivary gland weight. At concentrations which did not inhibit oviposition (20–50 g/kg b.w.) many of the eggs dried out even though they were kept at high RH. We then demonstrated inAmblyomma americanum, Dermacentor andersoni andD. albipictus that removal of egg wax (by extraction with hexane) induced a marked increase in water permeability. IVM neither increased water permeability ofD. andersoni eggs nor diminished the amount of egg wax deposited on the surface of the eggs, when injected posteriorly through the alloscutum. However, injection of IVM, dimethylsulphoxide (vehicle for IVM) or distilled water through the articulation between the capitulum and scutum (anterior injection), did markedly reduce the wax coating and increased egg permeability. We suggest that anterior injection damages Gené's organ and thus causes the latter effects.     

Wax lipid secretion and ultrastructural development in the egg-waxing (Gene's) organ in ixodid ticks

Gene's organ, the egg-waxing organ of ticks, performs an essential function in females by coating the eggs with a waterproofing layer during oviposition, which prevents desiccation of the embryo, ensuring its viability. The organ is a target for control agents and a potential site of virus replication involving trans-oval transmission of arboviruses. The organ is a complex dermal gland, developed to an elaborate degree. The external appendage, the horns, is an evertable balloon-like cuticular sac which manipulates the eggs and coats them in wax. Wax passes through pores in the cuticle from the internal, sub-cuticular lumen. Gene's organ develops in synchrony with oogenesis and oviposition. This paper describes the development of the gland cells and formation of the intra-cuticular lumen and its ultrastructure during engorgement and oviposition in ixodid ticks. The structural basis for wax secretion in Gene's organ is also described.     

Intercellular junctions in the hypodermis,salivary gland and gené's organ of the cattle tick,Boophilus microplus

Summary The intercellular junctions that occur in the hypodermis, Gené's organ, and the salivary glands of the tick, B. microplus, are described. The epithelial cells of the hypodermis are connected by spot desmosomes and septate junctions and the secretory cells of Gené's organ by septate and gap junctions. The cap cells in the alveoli of the salivary gland connect to adjacent cells by gap junctions, hemidesmosomes and septate junctions into which microtubules are inserted.The authors would like to thank Mr. R. Lamb for preparing the plates. M.W.J. Megaw was supported by an S.R.C. Studentship     

Experimental infection withRickettsiella phytoseiuli in adult femaleDermacentor reticulatus (Ixodidae): an electron microscopy study

Rickettsiella phytoseiuli naturally occurring inPhytoseiulus persimilis mites was cultivated in adult femaleDermacentor reticulatus ticks. It demonstrates all six known developmental stages: dense, intermediate, bacterial, giant, crystal-forming and small dark particles. These stages of rickettsiae were found in salivary glands, Malpighian tubules, synganglion, ovaries, tracheal complex, haemolymph, fat body and alimentary tract.Rickettsiella phytoseiuli did not infect the Gené's organ. It multiplied in female ticks in a manner similar to that in the typical host mite,P. persimilis.     

Modus operandi of oviposition in Dermacentor reticulatus (Acari: Ixodidae)

The process of oviposition in D. reticulatus was observed and found to be a sequence of exactly coordinated, interlocking events independent of the phase of oviposition. The average period of oviposition in the investigated ticks was 31.6 days at 20 °C and 95% relative humidity. The number of eggs deposited on each day increased until reaching a maximum on the fifth day of oviposition and then decreased continuously. As a result, most of the eggs were deposited during the initial phase of oviposition. The total number of eggs was proportional to the ticks' weight replenishment. Egg-laying commenced with the lowering of the capitulum and the simultaneous spread of the pedipalps which were lowered to the body wall embracing the genital aperture on both sides. Immediately afterwards the cuticular sac of Gene's organ was pushed out and retracted several times. At the cuticular sac's maximum extension, the vestibulum vaginae prolapsed, forming the ovipositor as an extended tube which handed over an egg to the two horns of the cuticular sac after a brief, but intensive, contact with the cuticular sac. Then the vestibulum vaginae invaginated, the pedipalps closed, and the cuticular sac was retracted. Finally, the egg was transported onto the dorsal area of the tick by means of a vigorous rising of the capitulum. During the course of oviposition most of the events, especially the period of egg embracement by the cuticular sac, were prolonged, as was the total time for laying one egg. Similarly, the intervals between successive egg-laying processes increased continuously.The number of eggs deposited was not dependent on the functional ability of Gene's organ, as shown by similar numbers of deposited eggs from ticks with and without mechanical blocking of the cuticular sac. But the participation of the organ in the process of oviposition proved to be a prerequisite for the viability of the eggs. Larvae developed and hatched only from those eggs which were deposited from ticks with an undisturbed Gene's organ. In comparison, eggs without contact to the cuticular sac of Gene's organ dried up and shrivelled immediately after being deposited and did not hatch. Consequently, it strongly suggests, together with the results from other studies, that Gene's organ covers the eggs with a secretion that prevents the loss of water.     

Novel diterpenoids and hydrocarbons in the Dufour gland of the ectoparasitoid Habrobracon hebetor (Say) (Hymenoptera: Braconidae)

Chemical constituents contained in the Dufour gland of the ectoparasitoid Habrobracon hebetor (Say) (Hymenoptera: Braconidae) were characterized. Three terpenes, beta-springene, a homo-beta-springene, and a homo-geranyllinalool constitute approximately 37% of the gland components, with the remaining 63% all being hydrocarbons. The hydrocarbons consist of a homologous series of n-alkanes (n-C21 to n-C31), a trace amount of 3-methyl C23, a homologous series of internally methyl-branched alkanes (11-methyl C23 to 13-methyl C35), one dimethylalkane (13,17-dimethyl C33), a homologous series of monoenes (C(25:1) to C(37:1)) with the double bonds located at Delta9, Delta13 and Delta15 for alkenes of carbon number 25 to 31 and at Delta13 and Delta15 for carbon numbers 33 to 37 and three homologous dienes in very low amounts with carbon numbers of 31, 32, and 33. The terpenoid and hydrocarbon composition of the Dufour gland was similar in virgin and mated females. However, in contrast to the hydrocarbons, the amount of beta-springene and homo-geranyllinalool increased significantly with time after adult emergence from the cocoon. Although many hydrocarbons in the Dufour gland are the same as those on the cuticle of this species [Howard and Baker, Arch. Insect Biochem. Physiol. 53:1-18 (2003)], substantial differences also occur. Of particular note is the chain length of alkenes and location of the double bonds: cuticular alkenes have a chain length of C23 to C29 and double bond locations at Delta5, Delta7, and Delta9, whereas the Dufour gland alkenes contains a greater range of carbon numbers and have no Delta5 or Delta7 alkenes. The Dufour gland contains only one of the long-chain dimethylalkanes found on the cuticle. Also, no terpenoids are found on the cuticle, and the Dufour gland contains none of the secondary wax esters that are major components on the cuticle. GC-MS analysis of lipids carried in the hemolymph of H. hebetor indicated that all hydrocarbons found on both the cuticle and in the Dufour gland are present, as are some of the wax esters. However, none of the terpenoids were detected in the hemolymph. This suggests that the hydrocarbons are synthesized in other tissues or cells, probably by oenocytes, and differentially partitioned between the cuticle and the Dufour gland. The terpenoids are most likely synthesized within the Dufour gland. Analysis of surface lipids from eggs laid within 18 h indicated that no diterpenoids were present. Rather, the lipids present on the eggs were n-alkanes, monomethylalkanes, alkenes, and secondary alcohol wax esters. This composition did not reflect that of the Dufour gland, hence eggs are not being coated with Dufour gland components during oviposition.     

Ultrastructure of the accessory glands of gene's organ in the cattle tick, Boophilus

The organization and ultrastructure of the accessory glands of the cattle tick, Boophilus microplus, are described. The glands consist of two groups of acinar cells situated on either side of Gene's organ. A single acinus consists of from eight to 12 cells and each cell is connected via an individual duct to pores on the dorsal surface of the mouthparts. The position of these pores is such that the secretion of the accessory glands is incorporated into the egg wax during oviposition. Each gland cell has striking quantities of smooth endoplasmic reticulum and numerous Golgi dictyosomes and appears to produce a secretion that is lipoidal in nature. Each cell secretes into its own individual lumen and is connected to a cuticular pore by a duct cell.     

Comparative reproduction and nonparasitic development of Boophilus microplus and hybridized Boophilus ticks (Acari: Ixodidae) under natural field conditions in subtropical South Texas

The reproductive biology and nonparasitic development of Boophilus microplus (Canestrini) and hybridized Boophilus ticks (B. annulatus (Say) male x B. microplus female) held under natural field conditions in south Texas throughout the year were compared. Comparisons between the two types of ticks indicated that the ovipositional biology (percentage of ovipositing females and number of eggs laid) of the females favored hybrid ticks during some months and B. microplus ticks during other months. However, on a yearlong basis, there was virtually no difference in the percentage of ovipositing females or in the number of eggs deposited by hybrid females as compared to B. microplus. The duration of each nonparasitic development period (preoviposition period, incubation period of eggs, and larval longevity) showed that both types of ticks had very similar developmental and survival rates during the year. Generally the difference in duration of each of the nonparasitic parameters was <8 days, leading to a high degree of synchrony of the nonparasitic developmental rates between the two types of ticks. On the other hand, egg hatchability of hybrid ticks was consistently lower than pure-strain B. microplus throughout the year with significantly lower hatch rates occuring in April, July, August, October, and November. Thus, results obtained on percentage of ovipositing females, number of eggs laid, preoviposition period, incubation period of eggs, and larval longevity provide positive encouragement for the possible use of sterile hybrid males as a means of eliminating a native B. microplus population. In contrast, results of the egg hatchability of the two types of ticks indicate that the selective advantage afforded toB. microplus could have an adverse affect on the success of a sterile hybrid male program by making the number of hybrid ticks necessary to eliminate a native population prohibitively high.This paper reports the results of research only. Mention of a commercial or a proprietary product in this paper does not constitute an endorsement by the US Department of Agriculture.     

Antiviral effect of the egg wax of Amblyomma cajennense (Acari: Ixodidae)

The control of viral infections, especially those caused by influenza viruses, is of great interest in Public Health. Bio prospection has shown the presence of active principles in the hemolymph of arthropods, and in the salivary gland of ticks, and some of these are of interest for the development of new pharmacological drugs. Ticks lay their eggs in the environment, and to protect them from desiccation and microbial attack they involve the eggs in a waxy layer produced by an organ known as Gené’s Organ. In this study, the eggs wax from tick Amblyomma cajennense (Fabricius) was extracted using ice cold phosphate buffer. The antiviral activity was evaluated with picornavirus and influenza virus. In both cases egg wax was able to inhibit virus replication. For influenza virus, an amount as small as 12 μg/mL of crude egg wax suspension neutralized 128 UHA (hemaglutinant unit) of H₁N₁ influenza virus. With picornavirus, egg wax led to a 256-fold reduction in virus production by L929 cells. Egg wax was not cytotoxic to VERO, MDCK and L929 cell, being observed that the cell morphology was preserved with concentration as high as 2 mg/mL. In addition no genotoxic effect was observed for Vero cells, suggesting a very interesting potential antiviral activity.     

The behaviour of laying workers and the morphology and viability of their eggs in Melipona bicolor bicolor

Abstract. As in many other stingless bees, Melipona bicolor bicolor Lepeletier (Apidae: Meliponinae) workers lay two morphologically distinct types of eggs: slender ones that have a typical patterned chorion, and larger ones that lack this pattern. In this paper we report on the relation between egg morphology and the behaviour of the workers that lay such eggs. In most cases, the laying of each of these egg types is accompanied by a unique sequence of behaviours. After a worker has laid the unpatterned type of egg, she generally leaves the cell, giving the queen the possibility of eating this egg. In the case of the patterned egg type, the worker usually closes the cell immediately after her egg laying. When worker egg laying occurs right after a series of regurgitations, it stops the queen from ovipositing. When, instead, a worker lays an egg after queen oviposition, the cell contains two eggs. This study also revealed cases in which workers laid slender, patterned eggs without closing the cell, and other cases where workers laid large, spherical, unpatterned eggs and instantly closed the cell. Experiments in which worker eggs, destined to be eaten by the queen, were protected by covering the cell artificially with a piece of wax showed that some of these eggs developed into larvae, although they were occasionally relatively small. The occurrence of a range of combinations of egg-laying behaviours and egg morphologies in M. b. bicolor workers is discussed from the perspective of worker egg laying in other stingless bees.     

Source of the host marking pheromone in the egg parasitoid Trissolcus basalis (Hymenoptera: Scelionidae)

After oviposition, Trissolcus basalis females always mark the host's surface, depositing host marking substances for herself and to warn other ovipositing females. The perception of these host marking substances, probably through the antennae, can induce the female to leave and seek healthy hosts. Parasitoid females exposed to conspecific parasitized egg masses left the host egg masses significantly more often than when exposed to non-parasitized egg masses. More egg mass leaving behavior also was observed when the egg masses were treated with Dufour's gland secretion but not when treated with secretion from the common oviducts. The common oviduct has a secretory epithelium that produces electron-dense vesicles, probably containing proteinaceous substances. The secretory cells of the accessory gland, Dufour's gland, contain electron-lucid vesicles, whose secretion appears to be a lipid similarly to that found in pheromone secreting glands. Ultrastructural and behavioral evidence suggests that Dufour's gland is the host marking pheromone source.     

γ-Aminobutyric acid (GABA) and other amino acids in tissues of the tick,Amblyomma hebraeum (Acari: Ixodidae) throughout the feeding and reproductive periods

We assayed a variety of tick (Amblyomma hebraeum Koch; Acari, Ixodidae) tissues for a number of amino acids throughout the feeding and early reproductive periods. Our HPLC assay could detect as little as 2–5 pmol per sample of the following: GABA, glycine, serine, glutamine, alanine, taurine, glutamate and aspartate. All of these amino acids could be detected in the salivary gland, synganglion (=total CNS in acarines), haemolymph, Gené's organ, seminal receptacle and ovary. GABA reached high levels in the salivary gland of freshly engorged ticks (685 nmol g^-1) and in the synganglion it exceeded 1000 nmol g^-1 throughout most of the feeding cycle and the first week post-engorgement. GABA also reached a peak titre in the haemolymph of 40 nmol ml^-1. Taurine levels peaked at 1065 nmol g^-1 in the salivary gland from large partially fed ticks. Glutamate and aspartate were likewise found in the salivary gland and synganglion at high concentrations. For most of the amino acids there is insufficient information to correlate these titres (and fluctuations of titres) to neuromodulatory functions. It is possible, however, that the high GABA titre in the salivary gland of engorged ticks is correlated with an augmented level of fluid secretion.Deceased: Department of Pharmacology, University of Alberta, Edmonton, Alberta, Canada, T6G 2E9.     

Relations of intraspecific variations in fecundity, clutch size, and oviposition frequency to the body size in three species of stoneflies, Sweltsa sp., Isoperla aizuana, and Stavsolus japonicus

Most stoneflies oviposit several times during their adult stage. In this study, the relations among oviposition frequency, the number of eggs per egg mass, and body size were examined in the chloroperlid stonefly Sweltsa sp. and two perlodid stoneflies, Isoperla aizuana and Stavsolus japonicus. It was found that larger individuals tended to oviposit more frequently than smaller ones, but the relation was significant only in Isoperla aizuana. In Sweltsa sp. and Isoperla aizuana, the number of eggs per egg mass was higher for the less-frequently ovipositing individuals than for those ovipositing more frequently, and the number of eggs per egg mass decreased with successive oviposition events. Stavsolus japonicus showed the same tendency but it was not statistically significant.     

The effect of egg load on readiness to accept a low-quality host plant is weak and age dependent in a geometrid moth

Abstract.  1. Most empirical and theoretical studies treat egg load as a major determinant of insect oviposition behaviour, and predict a positive link between egg load and readiness to oviposit.
2. In the present study, the correlation between egg load and readiness to lay eggs on a low-ranked host was examined in wild-caught moths, Scotopteryx chenopodiata L. (Lepidoptera: Geometridae).
3. No overall effect of egg load on oviposition could be detected. However, there was a significant interaction between egg load and wing wear: in only one wing-wear class, out of four, high egg load tended to promote oviposition, while in the three other classes the effect was insignificant.
4. The results suggest that the effect of egg load on oviposition may depend on the age of the ovipositing insect.     

Morphological evidence that salivary gland degeneration in the American dog tick,Dermacentor variabilis (Say), involves programmed cell death

During the preoviposition and oviposition periods of ixodid ticks, the salivary glands degenerate. It is unclear whether this is a necrotic or a programmed cell death event. We used an in situ TUNEL technique to determine if salivary gland degeneration involves apoptosis. Salivary glands were dissected from replete females at days 3, 5, 8, 11, 13, and 33 post-detachment. There were no differences in tick weight at detachment, suggesting that changes were not due to engorgement abnormalities. The onset of apoptosis began at day 5 and continued through oviposition at day 33. The greatest amount of nuclei containing fragmented DNA was observed on day 8 post-detachment, suggesting this was the peak occurrence of programmed cell death. Further, the temporal organization of programmed cell death suggests that the granule-secreting acini undergo apoptosis first, and that during the first week of oviposition the type I acini do not exhibit programmed cell death. These data suggest that the type I acini may still function in maintaining off-host hydration state of ovipositing females. These data provide morphological evidence that salivary gland degeneration involves a temporal programmed cell death event.     

Manifestations of immunity in sheep repeatedly infested with Amblyomma americanum ticks

Three sheep were infested 4 times with 100 Amblyomma americanum tick pairs and kept indoors until the natural termination of the infestations. Characteristics of the tick populations that show efficiency of feeding, fertility, and offspring development, and ELISA antibodies to tick salivary gland extracts were studied at each infestation. On average, female ticks fed on tick-naive sheep detached at 12.1 +/- 0.2 (mean +/- standard error) days, weighed 492 +/- 16.8 mg, engorged 40.7 mg per day, and 35% survived to detachment during the first infestation. During the fourth infestation, they detached at 17.3 +/- 0.8 days, weighed 321 +/- 14.4 mg, engorged 18.8 mg per day on average, and 23% survived to detachment. On average, oviposition of female ticks fed on tick-naive sheep started at 11.8 +/- 0.6 days of detachment, the egg mass weighed 236 +/- 13.2 mg, 43% of the female weight turned into eggs, and 89% of the ticks that detached survived to oviposition during the first infestation. During the fourth infestation, oviposition started at 15.1 +/- 0.5 days, the eggs weighed 103 +/- 9.9 mg, 13% of the tick weight became eggs, and 67% of the ticks survived to oviposition. On average, eclosion started at 35.4 +/- 0.9 days of oviposition and 83% of the egg batches hatched in the first infestation. During the fourth infestation, eclosion started on day 34.9 +/- 0.7, and 47% of the egg batches hatched. Anti-tick resistance was expressed as an inhibition of feeding, fertility, and offspring development.(ABSTRACT TRUNCATED AT 250 WORDS)