Regeneration of Haller's sensory organ in the tick, Ixodes rubicundus (Acari: Ixodidae)

A study of regeneration in nymphs and adults of the South African tick Ixodes (Afrixodes) rubicundus, from which the forelegs had been amputated during the previous instar, revealed that the structural changes in regenerated Haller's sensory organs resemble those observed in other ixodid ticks, in particular in another prostriate tick, Ixodes (Ixodes) ricinus. The adult regenerates re-establish their atavistic features in terms of the increased number of different sensilla on the distal knoll, in the anterior pit and the capsule. The nymphal regenerates, in contrast, re-establish the features of the previous instar through a reduction in the number of some sensilla on the distal knoll and in the post-capsular area. The structural changes in different compartments of the organ appear independent. The phenomenon of regenerative induction through the appearance of specific changes in Haller's organ of the contralateral non-treated foreleg is probably characteristic only of prostriate ticks. A unique modification in the regenerated Haller's organ as revealed by duplication of the Haller's organ capsule was discovered in both I. rubicundus nymphs and adult ticks.     

革蜱成虫哈氏器的扫描电镜观察

本文报道了用扫描电镜观察革蜱属8种成虫的哈氏器.其中三种成虫(阿坝革蜱D. ahaensis、金泽革蜱D. auratus和中华革蜱D. sinicus)的哈氏器结构是首次报道,并与其他属6种成虫的哈氏器进行了比较,以阐明其形态特征及其在分类中的意义.另外,还将这14种成虫哈氏器与各自未成熟期的哈氏器进行了比较.分析和讨论了同种不同时期哈氏器结构的异同.东洋区的金泽革蜱和古北区的革蜱种类间,在孔毛形状,位置及基盘数目上有明显差别,而后者中,边缘革蜱D. marginatus的孔毛位于近端外侧,草原革蜱D. nuttalli和网纹革蜱D. reticulatus的孔毛位于远瑞外侧,其余种类无明显差别.     

Ultrastructural studies of Haller's organ in the argasid tick, Argas tridentatus (Argasidae)]

Haller's organ in A. tridentatus consists of a capsule and an anterior group of sensilla. The capsule is the hollow in the cuticle on the dorsal surface of the first tarsus, where 4 pored hairs of olfactory sensilla are situated under the cover of the roof, formed by an anostomosis of the upper brunches of pleomorphs (capsule's bottom non-sensory cuticular outgrowths). The canal of the accessory ampullaceous sensillum opens in a capsule near the bottom. The anterior group of sensilla consists of two parts: proximal part, containing pored grooved and thin hairs, is homologous to the anterior grouf of ixodid ticks, and distal one which has no homologues in ixodids. Fine structure of all the sensilla in the mentioned parts of Haller's organ is described in detail.     

Regeneration of Haller's sensory organ in two species of hard ticks of the genus Haemaphysalis (Acari: Ixodidae)

A study of Haller's organ regeneration in nymphs and adults of Haemaphysalis turturis and parthenogenetic females of H. longicornis, from which the forelegs had been amputated during the previous instar, revealed structural changes in regenerated organs. The adult regenerates reestablished atavistic structural features, while the nymphal regenerates retained larval features, which is typical of regenerates of two other genera examined previously (Ixodes and Hyalomma). Data on regeneration of Haller's sensory organ testify to an ancient character of the genus, standing closely to the base of the phylogenetic tree of hard ticks.     

Fine structure of tarsal sensilla in the taiga tick Ixodes persulcatus (Ixodinae)

The first tarsal segment of the taiga tick bears 4 general types of sensilla (except for sensilla forming the Haller's sensory organ on the dorsal surface of the tarsus): tactile mechanoreceptor sensilla of 3 types, contact chemo-mechanoreceptor sensilla of 2 types, and double-walled pore sensilla. One of these types, the chemo-mechanoreceptor upper-pore sensilla, was found only in the Ixodinae, and not found in the examined representatives of the Amblyomminae. This type of sensilla was also found in the palpal receptor organ of the ixodid and argasid ticks.     

Bat ticks of the genus Argas (Ixodoidea: Argasidae). 13. Haller's organ roof and anterior pit setae in Carios and Chiropterargas

Adults of 4 of the 6 species constituting the subgenus Carios and of 3 of the 4 species constituting the subgenus Chiropterargas were studied by scanning electron microscopy. All species parasitize Old World cave-dwelling insectivorous bats (Microchiroptera). The anterior pit setae number 10 in Carios and 10 or 11 in Chiropterargas. In most Carios, the setiform seta is replaced by a second serrate seta. In 2 of the 3 studied Chiropterargas species, 1 of the 2 grooved setae is exceptionally long. Porose setae number 3 in Carios and 3 or 4 in Chiropterargas. The Haller's organ roof in both subgenera is solid, lacking perforations; the aperture is narrowly transverse in Carios, irregularly wide or wide and transverse in Chiropterargas; uniquely, 1 or 2 sensilla protrude from the aperture of Chiropterargas species. The protruding sensilla and long grooved seta of Chiropterargas suggest a probably distinctive sensory-behavior pattern common to these ticks. Other morphological characters are discussed and compared to show relationships between these 2 subgenera and the subgenera Argas and Persicargas and distinctive characters present only in adult and/or larval Carios and Chiropterargas.     

Effect of of methoprene and retinoic acid on characteristics of Haller's organ regeneration in the tick Haemaphysalis longicornis (Acari, Ixodidae)

The results of experiments on regeneration of the Haller's sensory organ in the metastriate ixodid tick Haemaphysalis longicornis Neumann in the presence of the juvenoid methoprene and retinoic acid and of studies of its structural changes by SEM confirmed the similarity of prostriate and metastriate ixodids as concerns the juvenalizing effect of the above morphogens on regenerative processes during nymphal-imaginal metamorphosis. However, the metastriate ticks (Haemaphysalis and earlier studied Hyalomma) are well behind the prostriate ticks (Ixodes) as concerns the extent of changes induced by juvenoids and retinoids in the sensillar sets of Haller's organ regenerates.     

Specific Features of Haller's Organ Regeneration in the Tick Haemaphysalis longicornis (Acari,Ixodidae) in the Presence of Methoprene and Retinoic Acid

The results of experiments on regeneration of the Haller's sensory organ in the metastriate ixodid tick Haemaphysalis longicornisNeumann in the presence of the juvenoid methoprene and retinoic acid and of studies of its structural changes by SEM confirmed the similarity of prostriate and metastriate ixodids as concerns the juvenalizing effect of the above morphogens on regenerative processes during nymphal-imaginal metamorphosis. However, the metastriate ticks (Haemaphysalis and earlier studied Hyalomma) are well behind the prostriate ticks (Ixodes) as concerns the extent of changes induced by juvenoids and retinoids in the sensillar sets of Haller's organ regenerates.     

Limb regeneration and apolysis in the tick, Ornithodoros tartakovskyi.

Limb regeneration potential and the apolysis process were investigated in the argasid tick, Ornithodoros tartakovskyi. Developmental instars received single or multiple amputations and were subsequently allowed to undergo single or multiple apolyses. Amputated ticks regenerated complete normal limbs but only after four successive apolyses. Following a single apolysis, the majority of regenerated limbs were essentially miniature duplicates of normal legs but commonly lacked normal chaetotaxy and/or tarsal hump(s). The site of amputation distal to the coxa-trochanter joint, number of limbs removed from an individual, and instar amputated did not consistently influence the extent of regeneration. Coagulation and clot formation were observed.The limbs of the tick apolysed within the old leg hulls. Larvae and nymphs amputated relatively early during the period of apolyses regenerated limbs; late amputations precluded regeneration. The process of apolysis was irreversible and not obviously affected by amputations.     

Perception of breath components by the tropical bont tick,Amblyomma variegatum Fabricius (Ixodidae)

Summary Wall-pore sensilla in the capsule of Haller's organ on foreleg tarsi of the tick, Amblyomma variegatum, show multicellular responses upon stimulation with human and bovine breath. Filtering breath through charcoal removes the stimulant for some of these receptors. Analysis by gas chromatography coupled with olfactory sensillum electrophysiological recordings indicates that an ethanol extract of the breath components trapped on charcoal contains a major stimulant eluting at the same retention time as H₂S. Two types of H₂S-sensitive receptors have been identified. They are housed in separate sensilla, and are called sulfide-receptor 1 and 2.Although, both receptor types are characterized by a high sensitivity to H₂S with an estimated threshold of ca. 0.1 ppb and a response range covering 5–6 log orders of magnitude, their overall response to sulfides and mercaptans is nevertheless dissimilar. The type 1 receptor fires slightly more upon stimulations with H₂S than type 2, whereas ethylmercaptan induces a stronger response from type 2, and dimethyl sulfide activates only receptor 2.In a bioassay, H₂S tested at concentrations of ca. 0.02 ppm and 1 ppm equally arouses 60% of resting ticks. Two-thirds of these ticks quest the air with their first pair of legs, and the remainder start active search. By contrast, H₂S at ca. l ppm in a mixture with CO₂ severely diminishes the locomotor stimulating effect of CO₂.     

革螨跗感器的结构和功能

本文进一步研究了:④厩真厉螨截肢前后的爬行行为,表明第1对足其感觉功能;②用古拉广厉螨分别截各对足驱避反应的对比试验,见到只有当截去第1对足跗节时失去嗅觉功能,而截去第Ⅱ、Ⅲ、Ⅳ对足时,各组都仍有嗅觉功能;⑤对格氏血厉螨、厩真厉螨、毒厉螨和鼠颚毛厉螨进行涂漆前后的驱避试验,显示当跗感器被涂满封闭,则嗅觉功能消失;④以0.5%结晶紫或龙胆紫液染色的截肢标本,观察了厩真厉螨、毒厉螨、格氏血厉螨、古拉广厉螨及尾足螨股一种螨,足Ⅰ跗节末端凹窝中,至少都存在两类感毛,钝钉型感毛和长而尖的刚毛型感毛;⑤厩真厉螨雌螨和幼螨跗感器的钝钉毛分别为8根和5根,另外各有2根短而尖的毛,分别测定了长度,描述了形态特点;⑥透射电镜观察厩真厉螨、毒厉螨等的跗感器钝钉毛,毛外围有表皮壁,壁上有很多微孔,内有中心腔,腔内有树突。属化感器——嗅觉器;⑦电生理技术研究,当用氨和醋酸的气体刺激厩真厉螨、毒厉螨的离体足Ⅰ时,均产生明显的应激电位差,充分证明足Ⅰ辩节有嗅觉功能。     

Morphology and structural organization of Haller's organ during postembryonic development ofArgas (Persicargas) walkerae (Ixodoidea: Argasidae)

Scanning-electron-microscopic investigations of Haller's organ in larvae, nymphs I, II, III and IV, and male and female adultArgas (Persicargas) walkerae ticks showed that morphology and structural organization change during postembryonic development. Stage-dependent differences existed regarding setal numbers of the anterior pit as well as formation and reticulation of cuticular projections in the capsule cavity. The anterior pit increased in size in the course of postembryonic development. It contained only seven setae in larvae, one conical, setiform and grooved seta each as well as two porose and fine setae. Nymphs I, II, III and IV and adult ticks had equal numbers of setae; however, one additional unilaterally serrate and grooved seta each were present. Setal length increased continuously during postembryonic development and attained maximum values in adult ticks. The capsule consisted of roof and cavity and was located distinctly lateral in larvae, slightly lateral in nymphs I and II, and in all other stages directly on the longitudinal axis of tarsus. The capsule roof showed a reticular structure. The slit-like main aperture was located peripherally and arranged transversally to the longitudinal axis of tarsus I in larvae. Nymphs and adult ticks had a central, circular main aperture. Stage-dependent cuticular projections of varying form protruded into the capsule cavity. Larvae had only single, free-standing projections which ramified slightly and communicated with each other. Projections were more heavily reticulated in nymphs I and II. In nymphs III and IV as well as male and female adult ticks, a long centrally situated tube of reticular appearance was seen, which was supported by a large number of radially organized and interlocking pillars and communicated with the capsule roof. In all tick stages there were always four porose setae present, arranged on the capsule floor.     

孤雌生殖长角血蜱的哈氏器超微结构与发育

为阐明长角血蜱Haemaphysalis longicornis孤雌生殖种群的哈氏器结构及发育特征, 用扫描电镜对其各虫期哈氏器进行了观察, 分析了血餐对哈氏器发育的影响。结果表明： 该种群幼蜱、 若蜱和成蜱哈氏器形态结构基本相同, 均由前窝和后囊构成。幼蜱前窝感毛6根, 位于同一基盘; 若蜱和成蜱哈氏器相似, 前窝感毛7根, 其中1根孔毛位于外侧基盘, 另6根感毛位于内侧基盘。各虫期饱血后哈氏器大小均比饥饿状态下显著增大（P<0.05）。幼蜱前窝与后囊面积比值与若蜱相比无显著差异（P>0.05）, 若蜱前窝与后囊面积比值与成蜱相比差异显著（P<0.05）。各虫期哈氏器均在发育, 且血餐对哈氏器发育有重要影响。幼蜱至若蜱期哈氏器前窝与后囊的发育速度相似, 若蜱至成蜱期哈氏器前窝发育快于后囊。本研究结果在一定程度上揭示了孤雌生殖长角血蜱的哈氏器发育规律。     

中华真地鳖断足再生对发育的影响

研究中华真地鳖EupolyphagasinensisWalker的断足时间及数量对发育历期的影响,以及不同龄期断足处理后发育历期间的差异。以8龄若虫为研究对象,发现其再生临界期为20~26日龄。不同日龄的8龄若虫断去1条后足,蜕1次皮后能再生的若虫从断足至蜕皮的平均时间与8龄的正常历期相近,整个8龄若虫的发育历期均比正常历期长,并随断足处理日龄的增加而逐渐增加。蜕1次皮后断足没有再生的绝大部分个体,8龄若虫的历期均比正常历期短。以3日龄的4龄若虫为处理对象,发现断去不同数量的足后(蜕1次皮后均能再生),从断足至蜕皮的历期及4龄的历期,均随虫体断足数量的增加而增加,但断去1条或2条后足处理,从断足至蜕皮的历期间及断足处理后4龄历期与正常历期间的差异不显著。不同虫龄的若虫断去1条后足的腿节后1次蜕皮后能再生的大部分个体,从断足至蜕皮的时间比同龄若虫的正常历期稍长,所延长的时间相当于从龄初至断足处理的时间,而断足后1次蜕皮未再生的其余个体从断足到蜕皮的时间均很短。     

Phenol and lactone receptors in the distal sensilla of the Haller's organ in Ixodes ricinus ticks and their possible role in host perception

Extract of steer wool odor was found to excite olfactory receptor(s) in a wall-pore olfactory sensillum on the distal knoll of the Haller's organ. Three active volatile compounds were revealed in this odor by gas chromatography. Electrophysiological experiments revealed two types of receptors (sensory neurons) within the sensilla examined. One type of receptor responded only to phenolic derivatives, such as o-chlorophenol, o-bromophenol, o-methylphenol, 2,6-dichlorophenol, 2,6-dibromophenol, 2,4,6-trichlorophenol, but not to o-nithrophenol, p-methylphenol, 2,5-dichlorophenol, 3,5-dichlorophenol, 2,6-dinithrophenol, 2,6-dimethylphenol, and pentachlorophenol. The other type of receptor responded only to gamma-valerolactone. It is assumed that these cells play an important role in perception of a host from long distances (10-15 m), which is typical of Ixodes ricinus ticks.     

Regeneration of Limbs and Sensory Organs in Ixodid Ticks (Acari,Ixodoidea, Ixodidae,and Argasidae)

We present a review of our own and literature data on reparative regeneration in ixodoid ticks (chelicerate arthropods). Ticks have a high potential for reparative regeneration and a close relationship between regeneration and development determined by similar hormonal regulatory mechanisms. These mechanisms depend on ecdysteroid hormones, which participate in the initiation of both processes, and juvenile hormones, which direct these processes either to the maintenance of larval features or to the development of nymphal and adult features. We present a detailed analysis of the regeneration of Haller's sensory organs in ixodid ticks and propose an hypothesis about the role of juvenile hormones in the modification of morphogenetic processes in this group. Furthermore, we present data on the effects of insect juvenile hormone analogs (methoprene and fenoxycarb) on the regeneration of Haller's organ, which support this hypothesis. Studies on reparative regeneration in arthropods provide a broader view of the problem of repair morphogenesis in animals.     

Regeneration of limbs and sensory organs in Ixodid ticks (Acari, Ixodeidea, Ixodidae and Argasidae)

We present a review of our own and literature data on reparative regeneration in ixodid ticks (chelicerate arthropods). Ticks have a high potential for reparative regeneration and a close relationship between regeneration and development determined by similar hormonal regulatory mechanisms. These mechanisms depend on ecdysteroid hormones, which participate in the initiation of both processes, and juvenile hormones, which direct these processes either to the maintenance of larval features or to the development of nymphal and adult features. We present a detailed analysis of the regeneration of Haller's sensory organs in ixodid ticks and propose an hypothesis about the role of juvenile hormones in the modification of morphogenetic processes in this group. Furthermore, we present data on the effects of insect juvenile hormone analogs (methoprene and fenoxycarb) on the regeneration of Haller's organ, which support this hypothesis. Studies on reparative regeneration in arthropods provide a broader view of the problem of repair morphogenesis in animals.     

Ultrastructure of Haller's organ in the tick Amblyomma americanum (L.)

Summary Haller's organ on the tarsus of the tick Amblyomma americanum (L.) (Acarina: Ixodidae; nymphal stage) was studied by scanning and transmission electron microscopy. It consists of a distal bristle group, (the anterior pit), and a proximal capsule which encloses several sensilla. The seven sensilla of the anterior pit (A1–A7) are all thick-walled and multi-innervated (2–9 neurons), but at least three different types can be differentiated. Sensilla A1 and A2 possess large, plugged pores (>1000 Å) and are the only sensilla with branching dendrites. A3 and A5 are characterized by a spoke-wheel arrangement of the cuticle wall and very fine pores (100–200 Å) penetrating the spokes centrally; A4, A6, and A7 do not exhibit any pore system but a single opening at the bristle tip is assumed.The capsule contains seven thin-walled, blunt-tipped sensilla, and several non-sensory cuticular projections (pleomorphs). All of these sensilla possess large plugged pores in the cuticle wall and numerous dendritic branches of several neurons (3–5) in the lumen. Glandular openings were found inside the capsule; their significance is discussed.The fine structure of Haller's organ supports the functions postulated by Lees (1948), namely olfaction for the capsule and humidity reception (among others) for the anterior pit.This research was supported in part by the Office of Naval Research, and by NIH Training grant ES 00069. Paper no. 3459 of the Journal Series of the North Carolina State University Agricultural Experiment Station, Raleigh.     

Sense organs in post-embryonic stages of Hyalomma marginatum marginatum Koch, 1844 (Acari: Ixodida: Ixodidae). I. Tarsal sensory system.

Light and scanning electron microscopic studies showed the differences in morphology and in size of Haller's organ in larvae, nymphs and adults (females and males) of Hyalomma marginatum marginatum Koch, 1844. The length of the anterior pit setae increases during post-embryonic development. The localization of these setae is the same in all stages. Six setae (one porose seta, two grooved setae, two fine setae, one conical seta) contain anterior pit of various developmental stages. In nymphs and adults more numerous pores appear on the wall surface of porose seta than in the larval stage. The structure of the capsule roof also differs in various developmental stages. Haller's organ of Hyalomma m. marginatum shows great degree of morphological development which is connected with the complicated life cycle and feeding behaviour of this tick species.     

The Haller's organ roof and anterior pit setae of Argas ticks (Ixodoidea: Argasidae). Subgenera Secretargas and Ogadenus

Adults of the Argas subgenera Secretargas (3 species) and Ogadenus (1 species) were studied by scanning electron microscopy. In each species, the anterior pit and Haller's organ are situated in a large dorsal hump of Tarsus I and the 9 setae of the anterior pit are characteristic of the genus Argas in structure and numbers. In A. (S.) transgariepinus, an Ethiopian-Palearctic crevice-dwelling parasite of bats, the Haller's organ capsule roof is solid with a slitlike transverse aperture. In A. (S.) hoogstraali and A. (S.) echinops, Malagasy soil-dwelling parasites of Oplurus (Varanidae) lizards and the hedgehog-tenrec (Insectivora: Tenrecidae), respectively, the Haller's organ is virtually unroofed but partially screened by arborescent dorsal projections from the posterior wall of the capsule, and the open capsule contains numerous fine pleomorphs. In A. (O.) brumpti, a soil-dwelling parasite of the hyrax (Procavia), other terrestrial mammals, and lizards in the Ethiopian Region, the capsule is also virtually unroofed and contains numerous fine pleomorphs. The unroofed capsule is probably phylogenetically primitive and occurs only in these 3 and 2 other Argas species. The soil microhabitat (in Argas confined to 3 of the 4 species recorded here), and the reptile or ancient mammal hosts of these 3 species, as well as the zoogeographical isolation of 2 of the species in the Malagasy Region, are distinctive in this genus of 56 species. The interrelationships between an unroofed Haller's organ capsule and unusual biological properties remain to be determined.