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SYUGANGLION HIISTOLOGY OF THE CAML TICK HYALOW4A DROMEDARI7(ACARI:IXODOIDFA:IXODIDAE)

BY

Aloya S. marzouk, Galila M. Khalil, Fatma S.A. Mohamedf,and Sohair Abdel-Kawy

U.S. NAVAL ME'-DICAL RESEARCH UNIT NO 3(CAIRO, A AB REPUBLIC OF EGYPT)

FPC NEW YORK 09527

8 9 3 0 7

THE 9OYPTit JOURANAL OF HISTOLOGY - VOL 9 - 160- f EOMMl "Is

* Egypt. J. Hiatol. 9 (2) 1986: 309-320.

SYNGANGLION HISTOLOGY OF THE CAMEL TICK HYALOMMADROMEDARII

fAWIr: Itiodoldea: Ixodla) *+

Aleya S. Marzouk, Gaillas M.. Khlai. Falma S.A. Mohtamed and Sohelr Abdel-Kawy

(From: Medical Zoology Depanrisnt, .WUied States (.aal Medicalf Researit ,Uri

Numbe'r 7he (NAU3RU-JL Cairo, andfrom Zimplogy Depwimnents. arulie. of Snene Ain Shants. Quata, ant

A L-Azhar Univervitis.

ABSTRACT tick-patholgn associations andof being the center ofN neuroendocrine activity, the structural and

-The syagamallom histology in femle H. functional organization of the nervous tissue of thisa~t~U Isaue mad eomamere wt other highly specialized arthropod group need tobe studiedticks. The aysilisman lying In a peul&I Iu to understand its bearing on the physiologicalhond dma l I '..by a ftand epra-supbsal activities of the tick.

pook sad alomne"a pa) a a adbmoldlgeal Thbe basic structural components of the fusedpot (d pars @1 Paea pewsa and the Coplax ganglionic mass. the synganglion. has beens tudiedopbthoaomatle gaugilot). The symaalom is histologically in the ixodids Doophilusemead ,1 a connective dame shesah. doe (VroboopMhn) alearams Dirula (Tu'nleneva, 1965).aaauliemma. The pedwuns fildasilas the L(U.)lerophCanestrii(itinjtondTtchell.

eelam ad Is followed by the eordeal meil.. 1973). Derma -eo, (D.) andmosad Stiles (Douglas.ab"e Ia separated fIm the ole by the 1943). D. (D.) piesm Hermann (Joffe. 1963), D. (D.)subpeuliourlim. Aaaoelation, motor mad vudbb Say (Obenchain. 1974). HaemmphymW.ean, eery cal types we the aoms vililel (H.) flava Neumann (Saito. 1960), Hyilomma (H.)laguther Wh 4 of 5 adI eel typs. compose ae debit= Schulze, H. (H.) saetoleem Kochcedeal -o of the ayugmgloa. Ushe &W~e eel (Tiien 15. (Hysasommaa), sgypiuustypeIsOdY disrlmled Isthe peerlrim.Filvelsb Shauif and Rhlleephebam (11.) smaalm Latreilleof ilabeam hoelmomal -e e said waredI (Satpa ct aL. 1971) and the argasis Arti.eunmedmesmewlate .eiophe. - (P deasa)azbwes aiser. Hoogasla nd Kobla

INTRODUCTION (Rosbdy and Marzouk, 1994). A. (P.) palema Oken(Robinson andl Davidson. 1914; Satpa et aL. 1971;

The tick central nervous system isone of the' Eisen et al., 1973). Orniabodores (Aleetoeota)suitable sites for multiplication of pathogenic heley Cooley A Kohis (Sonenshine. 1970). 0. (0.)microorganisms such as spirochetes. rickettsias and moubata Murray (Eichenbserger. 1970). 0.viruse (Bursdorfer. 1951; Aeschlinn. 1951; (PavIv~yam) parked (Pound and Oliver, 1982) andRothady. 196Z; Reinhart et aL, 1972; Diab et &L. 0. (0.) mevanyl (Christophers. 1906; Evans and1977; Zaher at aL. 1977). Solomon. 1977).

Bcauseothevitlimpoetaneofthsvsteminthe Recent investigations on Hyalomem (H.)

From Research project 3M t1t tO2BSIO.AD.242. Naval Medical Restardi and Devetopent Commad Naval MedicalCommand. National Capital Region. Reuhesda. Maryland. Suppted is part by coeract RR041-2"1O. NR20i.tt4t with the01(m~ at 'Naval Research. Microbiology prowrast. Naval Biology project. Aetinqtot. Virginia. The opinions and assentioncontained herein ate the private urs o1 the authors and aic not to hr construed as offical or as reflecting the views uS theDepartmnt of the Navy or at the tuvelat Svie &I larg.

+ Dedicated to late Dr. Harry "ootrau hn inifinted aod supported this wcork before he passed sway on 24 Pahrary 19st,

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deonsadari Koch have revealed hat this lick is This is an irregular layer of scattered glial Cells whichresponsible forth transmssion of severalIarhovtises lies beneath the neurilemina. The cells. 2.1-2.5 urn(Hiogareal. 197S; Wood et at.. 1962). thick. hame elongated nuclei, and ill-defined

boundaries (Figs. 1.4). The subpsernetwlims. "isHerein. we describe the syI5palhOf histology of layer of irregularly arranged glial cell&. ca. 3-3.5 urn

the female 11. deomd to provide essential tkig2 ethcrxfomhenupi.information for further investigations on tik(i.2."ae h otxfo h nuoie

neurosecretory ell distribution, and their possible The corlas, the cortex is composed primarily ofinvolvement in hormcse~mediated emins controlling neuronal cells or perikarya. and a few tattereial athe various physiological activities of the tick, cells (Fig. 1-6). Three major types of necuronal cells

are observed: the association, motor andMATERIAL and METHODS neurosecretory cell types. These cells are classified

Engorged femasle H. dr elil collected from according to shape. size, cytoplasm stainingcamels in the Imbaba market. Gizs Governorate. properties and distribution. Assoeatlan neurons.Egypt. were tsed to start a colony in the NAMRU-3 Twocell types. A and Bare included in this category.Medical Zoology laboratory. The ticks were fed on Type A cells from the bulk of the cortical layer. Theythe rabbit Orleteagus eusicetue using the method of are oval or rounded is shape, measuring 8.8- 10 um inflerger et al., (1971) at 25 ±10 C and 75% relative diameter, each with a central rounded nucleus.humidity. All animal care and manipulations were in measuring 5.5-6 urn in diameter (Figs. 1.3.4). Thecellaccordance with the Animal Welfare Act and nuclear boundaries are very distinct, theAmendment of 1976 (PLP 14-279 with subsequent cytoplasm is finely to coarsely granular. and theamendments). nuclear eltromautin is in the form of granular

inclusions. Type B cells, known as the globuli cells.For histological observations, the dorsal and occur in large numbers associated with the first pedal

sometinmes the ventral integument were remove4 and ganii and are the smallest neuronal cells (Fig. 3).the syngenglion was quickly fixed in modified They are, rounded in shape, measuring 6.0 urn inBourn's fluid (Humason. 1962) for 24 hours. The dimeter. with rounded relatively large nuclei,synglanglion. was then washed in 70% ethanol, measuring a. 4.5 urn in diameter. Cell and nucleardehydrated in anascendinj series ofethyl alcohol and boundaries are distinct; the cytoplasm is coarselydouble embedded in celloidin-paraffin. Serial granular and the nucleus is chromatin-rich.sectionss. 5-7 um thick, were stained by Mallory triplestain (MT) (Pastin. 1959). aldehydefuchuin (AF) Motor call. These are only few cells which are(Ewen. 1962) and the chrorme-hatmtoxylin phloxine easily recognized by their relatively large size (Figs. 3.(CHP) techniques (Gomori. 1941). 4). They are rounded to oval in shape. mearuing a.

16.5 urn in diameer. The cytoplasm is finelypranularand vacuolated arid ste nucleus is rounded in shape.RESULTS 7-7.5 emt in diaeter, and constains looser thread-like

The syngangion of femrale H. dromadai, lying in chrsomatis and one or two nucleoli.a periganglionic blood sinus. consists of a Nuoiceoycls hs el mdsrbtdisupraesophageal pan and subesophageal part, and in rserl toualy ciaerll Thsmeelarditrib ute inh

enclsedwithn awel-&fted onnctiv tinse cortex (Fig. 5). The eels are oval, rounded orsheath, the neurlemsa (Figs. 1)Lt The pinrineturiun irregu11111r in shaspe and vary greatly in size, rangingunderlies the neurilemma and is followed by the from lOX I1.5 to 55 x23.5um. Their nuclei am ovalcortical region, formed mainly of neuronal and non- or spherical in slope. nmuring ca. 7.7 X 5.5 urn toneutinet cells which is separated from the inner 11.2 X V. inn.fibrous core or neuropile by the subperineurnum.

Thes e lls contais mosecretory material (NSM)Thea ins This atath, 3-3.3 -m. is also in thefeormof floeculant veryfineorcoaegaiim.

reflected into the esophmgsal canal inside thes The stailaingaffintyofthiasaterisldfirnaeordngsysilanglion to surround the iesophagus as well (Fig to the stain usid. The NSM may accumulate at the7). In MT-Staned preparation, it sin blue Periphsery of the eel or around the ancleus and arenmsug s..in a collagnm =anire. The i mnm. sMessese ditribuuted throughout the cytoplasm.

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TauSWIemCUCo. C resnhjiofamifad H. among type A association eaits (A c. d. #list cells&inassar thatie the vascular sheath (v Sb) types c aud d; NI. betueilsrme; Pe, parintur (X

asociation cells; E, ovsophallos; 0C.giaiaek N.aamtal d NL, wuilsjuenn Np. nawopihe; Pa. (fig 5)

peiasiwot Spa. ahpauseasa (X 400) Froffial tttoa( ofe bsayuptallho of unfed (esata(FI&.3) H. itowemil thostal a neuscratoy cell INSCI

Tmrassesn tti of the sympigion of uniod (X 723,ktans H. dauI lis the tegion of the rust pedaljangla (P~e) aborwifig type 8 asaociation calls (3) (i.6and motorc alks (M). Letaning - is Ftgs. 1. 2. (X T 5ws a -etion f45*nalo of notad

723 L tensle IL drendar htowing different types of4 Fig. 4)glint calls (a. Isc. d. a). L. aophtallmer NI.

Traunsee seam.ct of the ayaglion of retailed Mailmes (X 4"'.

J12

Glht call. These cells iniclude 5 types differng in Ventral glonaendl. These two glomerutar pairs arelocation. shape, and ize. Typ a ese cells ame veistrotnedial to the protoccrebral neuropile I(Figs. 9.irrggularty distributed in the acuropilinand many of 12) and lateral to the stonsodeal polls. and arethem aecurcnhiate in the area adjacent to he boardered laterally by the bases of the cheliceral andesophageal croasn$ (Wig. 6). They are oval. pedipalpal neuroltikms The outer pair is larger (ca.meaaunng ca. 7 X 3.5 umn. and their oval nucei 18.7 uminiameter)than the inner one(ca. 13.2 ummneasusre ca. 5.5 X 2.2 umn. Type Ill. Thensellts occur in diensetes). Both pair are closely associated withamong the different neuronal cells (Fig. 6). They are the esophageal canal and stain darker than themome or Wia rounded in shape.nmeasuring 6.6.6 umin protocerebufal fleuropile. indicating a higherdiameter. and the large nucleus occupsies tMe majority neuropilar density.of theell body. Typ "e.Thsen cellsare scattered in The ehelleerial ug.This pair of ganglia

she eriherl nevesansngth neve iber(Fi.4- occupies the dorsolaterall part of the supraesophageal6). They afe densely packed and best observed where Part (Figs. g-10). Anteriorly. the neuropile appears asthe nerves commnunicate with the corresponding 2 sphrcal structures connected ventrolaterally with.europsde regions. They are rounded or oval in shape.measuring 4.3-6um indiameter. Typed ".Thenecells the anterior boarder of the pedlipalpal ganglia (Fig.16rM the peuineuriust beneath th OffkM ) Ventrally. the cheliceral neuropiles am interrupted(Figs. 1. 2. 4) They nwsr ca. 6.6 urn X II urn. across the midline by the protocerebrum.Type "e. These cells form the subperineurial layer The pedipailal pallia. This pair of ganglia is(Fig. 6). They are spindle to oval in shape. measuring posterior to the cheliceral gansglia. Aniteriorly. theyca. 9.9 tim X 3.5 umn. extend obliquely upards (Figs. 8. 9) and ace

connected together Posteriorly by a possesophagealTh~ee a ehal paf Th i pan i fomedas comnmissure (Fig. 9). The posterior aspect of both

result of the fusion of the protocerebrum, the pedipalpal ganglia aeadjacentto he anerior margin

cbeliceral ganglia, the pedipalpat anglia and the of the first pair of pedal ganglia (Figs. 10-12).

stomodeal pons(Fi. 7-12). Thepollaeeebrom.The The stamoideal POas. This it the most ventralprotocerebrnum appears as a single. median mass region of the supraesophageal pan (Figs. 7-9).which extenids in the dlorsovenrttal direction between Anterodoisally ad asirvstal.the stomodealdhe bases of the chelmeeral and pedipelpal neuroptils polns neuropile is boordered by a cortex formedand donal to the slatodeal poitsa(Fills. 8-42). The mainly of type Aassociation celtsand neurosrctor)proloccerebrnm contains the optic gonglia. discrete cells (Fig. 7). Posteriorly, the stomodeal pons isbiltesally-arnaneged glomeruli of hiagher neuropilar blardeved laterally by the pedipalpal neuropilesdensity and a number of interconanecting tracts (Fg. (Figs. 9,9).7. 9412). The glomecrsli include the anterodorsal.poeterodotul and ventral glomerui (Ftgs. 9-12). The subhesophageal part. This part includes theOptic Vin. This pair of ganglia occurs in the most neuropiles of the four pairs of pedal ganglia and theanterodlorsall par of the protocerebrum (Fig. 7) as complex opisthosomnatic ganglion (Figs. 12. 13).small. rounded neuropiles with their base coinciding Th "I prilla. These 4 pairs of ganglia arewith the anterior boarder of the eheliceiral ganglia. nietamerically arranged, the first being anterolateral.The oullular cortex surrounds; the lateral and ventral the second and third occupying the mnidlatersl region

aspctsOf hisganlia An enoesl ~and the fourth oriented positerollateral in a dorsalThese ae two glotnslar pain in the middorsal direction (Fig. 12).mein of the protocerebeal neuropile (Fill 10). Theyare spheuical in shape, the outer pair (ca. 20.9um and The ainterolateral margin of the first pedal gangliaa. 14.3 um in diameter, respectively) is larger than coincides with the posterior aspect of the pedipalpalthe in one. A very thin doual cortical layer ganglia (Figs. 10-12), while the anterolateral marginsserats, these glonsensli from the pesmneunium. of t second. third and fourth pedalganglia coincide

Pealudne d g ee. This pair of glonseruli, with theposterior aspect of the proceeding pedalmeasuing 17.6 um n l diameter. is situated ganglia (Fig. 12). The posterior margin of the firstpoaerolaterail to theanserodonaul gioomli (Fig. I I) three pedal ganglia coincides with (he anterior aspect

313

COWN

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-* I

9 ~*

(Fig. 7-11)Transurom uedictoofim syoaglio ofsmld N.b..udeftohowisti the paaliosuc structure ofhe supraocophaipalpean (Sp P) muml subiiioptbap panl (Sb P). Ad GooamIiadorsa gJomnih.4 S. type Bassocitionfttdh; Chi G. chetierlpas*i L. onophwwm KI souulheumaOrK.*tuy know Op G.oplicpugla; P E C. poteoge alcomiurP G.. first peda poalm Nd G.. uueoral aloinesilk PS S, Pensomilioic sinus; Pp G. pedipaipui aglia: Pr.

* procevebeut St P. stamodeal pOOK v GM. wefilul glocruli (X 2SO0t

Fromoal im deb usynpmgicuned funkH.bduuUwhovuiqskami ni6f the fourpedaPqlia(PG,-J taad tin opiulsoomi postim (Ope. 0). L cesophosoms' Sts P. uuboao~kwl peat; Spi Pm upnoohal Pon; V

Ca. Ventrl oalomul (XIO)

314

of tdo bcloag pedal peoua all the poetrmial shapIed emuatfatr occurs directly postenor to theaspect of tme foath pedal peak coicdes with the esophei and extends between the Podiplaluammbletel malt On cth pisthcacuatic pnglicts sia. Mue third, tombt and fifth commtuurus are(FIg 12)L SMAildensle OnuWlab. th.e faetuY staight and counect the scond, third and fourthkihow and marrdcc .hbspbstiml wasrs formed of Pda P94l%. repec~tiey. The eXtreities Of thetype 0 aucinida, call are amacheed with the ftl sixth cOminur ame directed ulightly toward theped pops (Fil.t 9, l@l. pcWior Tegll= of the sygungion. tin seventh

ccm..aae surrouuds the opisthosorwatc gunglioltThe -k cgac-mde Po-. Thi eduy

complex gaglion is located between the iteirypotteromial borders of the fourth pedal VD&ui in The m i lel consists of 2 modenately-dente.theuoelposor mon ofmbohe apan(FWg bilaterally symnmetrical conoecime which extend12). The comnplex natue *(thisagion is indicated from the region behind the emophagus to Termnaaeby mutltiple dorml and ventral neuroprie projections. anterior tO the opisthosomitic pnghion (Fill. 15).The, cortical baye surroeding the neuropile is Thes coniecivea give off fibers to the pedal pgigliarelatively thick in the ventedl region, and is and ane connected anteriorly by a postesoiphagealinvagmed into the neuropile dorsally Livinlg it a cOiiiiistlft which give off fibers anterolaterallY totribbed apperanc (Fill. 13). the pplpal gungin

Cuiidhnua dc ecuuihlms The horizontal lbe thkd leael includes a dense. fibrous, medianconisctives and vertical cominnes of the europile ningfrom which radiate one pairof connectives to theoccur in five leels. The &W klees the roetdorsal Padip&Ipel PEBWi ad one pair to each of the fouroneit which 7 dislinguinhad coanmrscco, (Fg, Pain Of Pedal IPOn&i IFil. 16).14t. The anteriormn. arc-shaped comumureOccurs in the supIeSAophageal Pent and srrunds the Il ctill leael consists of 2 pairs of connecties inclophaem anteriorly. The other six commissurs ame the center of The synganglon (Fig. 17). One pair ispresent in the subesophigeal part. The second sft- dorsal medi ad straight, and the other pair is more

Ternserse section of she oithosotic ginglon lops Ga in senliled Ivroake H. diuseareil (X 2.50)

I kill,54)Frontal action of htenynnanglion of miled H. oadrliai thefisi level olconaseiinand commisonshorrsgin

* ~~comsninuse 5Mr 1-7). Ops G. spiulsowosatic fangion: PS,. lingtso fourthl pedal ganglia; Sp P. ssspaoesphaegu parttX 11.0)

*Frontal atio ose synpglon alud H - Ioa ilig. The oeod l ofcosiicti r and aommittrrs showing two

ciannectise (V) and one conissesa lMrt. Ops. 0. opistlsosomtic aimqton: PS, firmt to loaflh pedal ltanglia; Sbi P.ssabeeopuastat part; Sp P. supasopamnal paen (X 1(m)

te 16)Frontat section of the synaargion of unfed It. dunmedwil ai thte thtird level of connectiveand commiiore-towingsmedian rnest I from whtich radiates connectiwes I toathe pedipelpal ginglia and isniach o1 the loar perdal pnkglus (VII

Il, 11)F~rontal action of the Pysangilion of wifed feuale It. uIdaI at the fourth level at conncttive, and commisare

nsowieg two pain of connective; (I. V,) (X IMS)

* ~~Frontl netleof the syrenlnossssfd finle Mdcsalthlll ee lonciraacnn~s iwthret pain of connective. IV 1-3l sod threcominuen tMr 1-3). Of K. otactory knots (X f10)

__________________________________________________________________________________________-.- -,~---- -. V

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ventral and is formed of widely separated connective% raicanpim (Bsnnington and Tatchell. 1973). D.which gives off dorsoventral fibers touech of the four amideesend (Douglas. 1943), D. pletuaf offe., 1963). 0.pairs of pedal ganglia- vwdf (Oinchisin 1974j. Hieemptiysala Elars,

miamishaelconssofshme pain ofconectives (Saito, 1960). llydemsa detitim. H. anstolesliand 3 commissures (Fig. 19). The median pair of (Tirvilenes. 1965). H. aepetons. . stgodlessconnectives extends between the olfactory knots and (Satija et 8L. 1971) and the argassids A. arbocemthe opithosomatic ganglion. The other two ame (Roshdy and Marzouk. 19841. A. fatisdien( Robinsonlateraelly situated, adjacent to each other anid join the and [bvidsoui. 1914; Satija et al.. 1971; Eisen ei al..second, third arid fourth pedal ganglia of each side. 1973). 0. ballep ISonenshine. 1970). 0. motabstaThe 3 coromissuresextend between second. third and ( EDheberlr. 1970). 0. laurtel (Pound and Oliver.

fourt pedl gaglia.respctivly.l982)and 0. mvsl(Christophers 1906. Evansandfoufh pdal angia. espctivly.Solomon. 1977).

The Peripheral nerve trunks and esophagus in H.

DISCIUSSION dimiai sYOngglon mr surrounded by apengsanglionic: blood sinus, thus simulating other

The synglanglion structure in keial H.droudaril ixoditiiaid argaisid ticks and most arachnids (Bullockshows the main evolutionary charteritics Of the and Horridge. 1965; Babu, 1965; Marzouk. 1974;nervous system in the other acarines and in the Rosldy and Marzouk. 1994. Marzouk et al..chelicerala, in general (loffe. 1963. 1964; Tsvileneva. unpublished data). However, this sinus is lacking in1964. 1965; Satija et al., 1971; Binnington and the mesouligmasid mite. MacroceulesTatchell. 1973; Obenchnin. 1974). The cheticefal and *iNeaademestle (Coons arid Axtell. 1971) and thepedipalpal ganglia location laterad to the prossigmalid mite. Blankasartla acmsseatellarisprotocrembrum in the anterior region of the (Mitchell. 1964).supraetsophagesl pant and the anteroventnal positionof the stomodul Pons in this part are common The neurilemmas inHl.drissidariiisawell-difined,fuatures in the topography of the tick synganglion collagenous. connective tissue sheath closely(loffe, 1963. 1964; Tsvileneva, 1965; Eichenberger. enshething the nervous tissue and extending into the1970,. Obenchasin, 1974; Pound and Oliver. 1982; esophageal canal inside the synganglion to surroundRoshdy and Marzouk. 1 1114). As in other ticks, the the esophagus. Transmission electron microscopy ofoptic renters in ff. &roweeer protocerebrsm are the neural lamella in D. varlialif, Aniblyommasrepresented by small glomerular formnations, and the amelase and Argaswroeas (Coons etal., 1974)corpora pedunculata is represented by anierodorsal has demonstrated repeated layers of homogeneous.and posterjsdorsal glomteruli, lackiqgloknerularcells finely granular. disorganized material, showing(Hgnstrom, 1928; loffe. 1963; Tsvileneva. 1964; axons which contain netroecretory vesicles. AObenchein. 1974; Roshdly s&W Marzonk. 199). feltwork of collagenouis fibrila showing periodic

cram-banding and embedded in an amorphis matrixThe suhesoiphagpeal pant of H. dromdarll is found hetween these layers. This composite

synganglion isformed as aresutofthe consolidation structure of the tick netirilemma is structurallyof four pairs of pedal ganglia and the complex similar to insect neuiral lamellar sheath, which allowsopisthosonsatic "agion. The frst pedal ganglia are relative permeability to nutrients and ionic exchangesituated along the side of the esophagus, due to the (Hess. 1958; Gray. 19*6 Beccetti. 1% 1. Asshurst andforewardllmovement of the pedipalpsal ganglia into the Chapmnan, 1961; Smith. and Treherne. 1963;sispesoplapal Part (lo11c. 1963). Olfactory knots. Asasurst, 1965. 1910; Assastant and Costin. 1971;associated with the rust pedal nsiropile, nmy have a Tieherne and Piebon. 1972). The tick neuirilemmaespossible sensory relation to Halice& organ acts peimaly as a protective sheath providing(Tsvilanseva. 1964; Obesichain. 19741, structural support for the neural architecture, and

Histilogical examination of the female H. may hae a functional similarity to that of insects.beeiisynglaglions revealed basi structural The jiermneirium in H. deemeari is a relatively

COtiPOsents similar to dhsae previously described in thin leyer of gI cella that lies below the nestnlemsnm,t ixodds L. ealeauliat (Teviemava. 1905).3L thus resembling other tiks (Ekicheberger, 1970;

317

Obenehartn. 1974). However. in A. aiborms. the tracts in the subesophageal part of H. dsvsmedarlpennetunm foems a complex series of extensive, synganglion is similar to that described in I). pies.tortuousa fembranousiaetenion$ Of gliallcell wich (lofe. 1963). H. deigat=a and H. -an-oeVcontain large intraciellnlar vacuole, with narrow (Obsenchain. 1974) and A. arboreus (Roshdly and

.' inteeelbslar spaces between adjacent cell bounsdauies Marzouk. 1994). Pound and Oliver (19812) suggest(Coots et at.. 1974 A notable difference between that this system not only provides linkage of ganglialixodide and augaSids Occurs in the stutua elements occurring within each of the two majororganization of the penncuumal gSiat cells (Coons et gang1ia[ ports. bustaso integrates postage of impulsesal., 1974 X In the ixodids Amb eomma ainavlam. between them.3L ndmip andi D. walsbili lConns et al.. 1974, The present anatomical data on the cellular

Sioningtoss and Lane. 1960) these cells lack orniaonfthsyggin nepoidsbeiracellular spaces and contain large Slycogen linenidato tolrf the mehanism cofteneprovidcrebisnfdeposts. The intacellular spaces are thought by fn aat lrf h ehns fnuoerto

thos auhor torefecta dffeenc inthetrohic and hormone production in H. dromidarli. Work in

requrements in the two families ofticks. Iinnington thsdrcinsudewyand Lane (1991)) suggtsted that the increase in the ACKNOWLEDGMENTSglyciogrin level, n LeaIpI perincurial cello mayprovide an important store of energy for the central We are greatly indebted to Professor Mohamed A.nervous system as in the cockroach. poslplanasa Roshdy. Department of Zoology. Ain Shamsamericaas (Wileaworth. 1960). The University and Scientific Consuttant at Medicalsssbperinetrnussn layer of g&I cells separating the Zoology Department. NAMRU-3. Cairo. for hiscortical zone from the neuropile in IL I 1 encouragement anid valuable comments, and forsynganglion is similar to that described in tbe mie critically reading the manuscript. Thanks are also dueMoaieshl umacdaoougle (Coons and Axtll. to Dr. William H. Dees. Read. Medical Zoology197 1). and the ixodid tick D. wasahMM (Obenchain. Department, for his interest in this work.1974). It is not found in other ticks (Tsvileneva. 1964; REFERENCESEicbenberge. 1970).

1. Aesehfilmama. A. (19S$): DeveloppemnentThe dastribution of the neuronal cell types in she embryonnaire d*Oralthodoros moubata

cortical zone of H. dromedanllsyngenglion resembles (Murray) et transmission tansovarienne detaht described in other ticks (Eichenberger. 1970. Baela dusltood. Acts Trop. IS: 15-64.Coons et al.. 1974; Obenchain. 1974; Obenchasin and 2. Ashit DE. (1965?; The connective tissucoliver. 1975; Rosbdy and Marzouk. 1964). sheath of the locust nervous system: itsAssoiaton. motor and neurosecretory neurons in H. development in the embryo. Q. F.I. Microsc. Scidromedal usually occur in bilaterally symmetrical 160:61-73.If groups. As in other ticks, the first pedal ganglia in H. ~ A~snt L(96:Tecnetv iseodeomudaul contain small compact masses of type B 3.Aine.Anl. DReX ('nit The conet-e74 sueassociation neurons forming globuli cells. 4Aimcu. An.L Rev Ent .13457.A(1 )Th

Four of five types of gliat cells occurring in the connective-tissue sheath of the ner',ous system otcortex and neuropile of H. dromudiri synganglion Loctai migraoela: an electron microscope

re similarto those described in A.aerors(Roshidy study. Q. FJ. Microsc. Sci. 162: 463-467.ad Marzouk. 1984). D. piet. (loffe. 1963), H. S. Aasti, DX ad Castle. N.M. (1971)p Insectsapielaand R. sassguineus (Satija etlal., 1971)snd muccoubstances. [I. The mucosubstance of the0. socialat (Eichenberger. 1970). However, the fifth central nervous sy'stem. Histochem. J. 3: 297-3 10.type ("c" cells) is only distributed in the 6. NOWs K.S. 41965) Anatomy of the centralsubpenineuraal layer of H. dromedari and also in D. nervous system of arachnids. Ph.D. Thesis. Srivarlabik (Obenchain. 1974; Obenchain and Oliver. Venkatesivara Univ. Tirupeti. Andhra. Pradesh.1975. India ZoI. .9,. (Anat.). Bid. 62. 5: 1-154.

7. BaebukI &. (1961): Indagini comparative sullThe basic organization of connectives and ultrastrutturs della fibrilla collagene nei divers:

commissures in a five-level system of nerve fiber ordimi degli insetti. Redis 4: 1-7.

318

8.p w 111s.S4 0isati.C.asod Chbow.Y.S.41973): distribution of necurosecretory cells in the brain.Demst.oration of a sex pheromone in three (tderstepoofl. J. Vet. Res. 44: 127-130.species of bard ticks. J. Mod. Eat. &! 84-46. 21. Ewen. A.D. (19112N An improved aldehyde

9. Blag" K.C. sod Tatehoill. R.J. (129) The fuchbin staining technitue for neurosecretoryservous sysanIte sad cumioecretory cells of Products in insects. Am. Microbcopal. Soc.ooopf sideaph (Acana: IxOdidlae). Z. Trans. 91: 94-96.Wine. Zool.. S.A. IN: 193-206. 22. Gray, E.G. (2959) Electron microscopy of

I0. Oningo". K.C. an ae. N.J. (IMP) collauen-lIikeconnective tissue fibrils of an insect.Perineurial and glial cells in the tick loophils Proc. Rt. Soc. B. IN: 233-239.in r.l (Acarina: Inodidack freeze-fracture 23. Gomnori. G. f2941): 'l0bservatioms with

and trace. tuis. J. Neurocytology 9. 343-362. dilfraida Unaion on bisan lIdas of Lanigerhans.I I. owsias. T.H. sod Hoenidge. G.A. (1985): Am. J. Paillol. 27: 395-402

Structure and Functions in the A~ervousSystem of 24. Havotrhn, 2. (1929): Vcrgleichcndt Anatonmteinvertebrates. Son Francisco and London: W. H. des Nesvensystetns der wirbefloven Tiere unterFreeman and Company. 2 vole.. 2517 pp. besonderer Berllcksiclstigung seiner Funktion.

12. *uargilorfer, W. (1951)I: Analyse des Berlin Springer Verlag. pp. 1-628.infektiomtverlaufex bei Ouldhodouie inoulata 25. Huow A. OMS) The line structure otf nerve cells(Murray) und der saturlichen Ubertragung Von and fibres, neuroglia and sheaths of the ganhlionspiroclueta duttoni. Acta Trop. M! 193-262. chain in the cockroach (Penliplanet none Ieant).

23. Chrallteb &R. (19UX) The atnatomy and J. Biophys. Biochemvcytol. 4:731-742.histology of ticks. Sci. Mesn. Offrs. Med. Sanit. 26. Hoogatnawl. H. (191S): Book review (The ittodidDep. India, n.s., (23) 58 p-. ticks of Kenya). J. Parasit. 61: 4.

24. Coom. LB. sod Axtull, R.C. (19711k Cellular 27. Hosanas. G.L I 1%2N Animal tissue techniques.arpizalion in the syntlanglios of the mite W. H. Freeman and Co.. San Francisco. 468 pp.Macroeheles touneaeiomsstic (Acarina: 28. aloft I.D. (1963N The structure of the brain ofMascrochelidoey. An electron microscopic study. Dernascantor pieces Herm. (Chelicerata.Z. Zelifonsct. 119* 3094321). Acarina), Zool. Zh. (Moscow). 42: 1472-1494.f In

15. Cown. 1-.4 Polay, M.A. and Axtel, R.C. Russian),(1174): Fine strscture of the central nervous 29. loffe., 2.0. (1194): Nerve apparatus insystem of D"esacetor vaablla (Say). bernaceatfor pietuas Herm. (Chelicerata.Anailyonnma amelintmu IL.) and Aegna Acarina) and its secretory function (in relation tomirberus Kiser. Hoogntroal and Kohis diapeusec). Avtoref. Diss. Saisk. Uchen. Step.(lxodloidea). J. fiamenit. G&687-699. Easel. iol. Nauk (Moskov. Ord. Lenina Trud.

w6 0kb, F.M. mid Slesma. Z.O. (2977): An Kristin. Znam. Gos. Univ. im M.V. Lemonosov.experimental study of ", Nll angel. in 4 Biol. - Pochven. Fak): Moskva. 26 pp.specensolfla tick.Spi-cietelocalizationand 30. Naiars A.S. (1974): Anatomical anddemities. L. Parasitenk. S& 201222. histological studies of the nervous system in someI

27. Dialbs JRk. (2941) The internal anatomy of terretrial arthropods. M.Sc. Thesis. Ain ShamsDum sI . aiernow (Stiles). Univ. Calif. Univ.. Egypt.

Dubl. Ent. 7:207-272. 31, MIIlboL a. (964): The anatomy of an adultis. Eilballoilir, G. (1970) Das Zentralnerven- chimer mite Onabart aetmellarle tWalchi.

sse onOnabdonfa modat Murray. J. Morph. 1142 373-392.

Entiickluafg. Acta Tropi. 27: IS-53. anastomical relationshipsofthesynganiglion in the29. Kb.. Y4. Warburg. M.R. aod GOW. 3. 411973)c American dog tick. Dissnenor varlablis

Neuroateretory, activity as related to leeding and (Acari: Ixodidae). J. Morph. 142: 205-223.oollonesia in the fowl tick Arpaps freulOken). 33. Gbom~ab F.D. ad Oliver. J.H. (2975)Gens. Corop. Eadocrinfol. 21: 331-340. Neuroscretory system of the American dosgtick.

20. Evams, A.A. and Sallonrn. 5.3. (tHll) Duarnen metrll (Acari: Ixodidae). If.neuroneeretion in Ornitbodor"e savigayl Distribution of secretory cell types. axonal(Audouin) lzodoidca: Arjasidae. The pathways and putative neurohemal-

319

neurocadoerlfl aeaociat1ls comparative Aipa UIn Oken (Amarna Arachnida) Zoot.bistological ad amsemimi uilkict J- Pot. 21:197-221.Morph. 1d9; 2W293. 4Z Smids. D.S. and Tralliens. J.L (1943)r

3. Ptei, C.F.A. (390): Notts on mnicroscopical Functional aspects of the organization of thetechniques for zoologists. Cambridge University insec nervous system. In: Advances in InsectPoe.s, New York and London 77 pp. Physiology. Vol. 1. J.W.L keamat. J.F.

35. Po4.. Owd Olive, Jill. (3IMP) Sysganglial Treherne and Y.B. Wigglerworth ed. Academicand neurosecretory morphology of female Pre.s. London arid New York. pp. 401-494.

COnlhnodoas pairsllCooley (Acari Argasidae). 43. Soesn es, DX (3976): A contribution to theJ. Morphology 173: 1W9-77. internal anatomy and histuiloggiy of the bat tick

*36. Ratakard. C.. Asscillns A. and Hacker. H. Owlthodores kelayiCooieyand Kohls. 1941.1I1.(1972): Distribution of Rickettsia-like The reproductive, muscular respiratory,microorganisms in various organs of excretory, snd nervous system. J. Wed. Ent. 7:

*Orailhodoros tonaie laboratory strain 289-312.lxodoidea. Argasidae) as revealed by electron 44. T sedte, J.E. and Pichen. Y. (1972): Insect

microscopy. Z. Parasitenk. 30.201-209. blood barrier. Adv. Insect Physiol. 9- 257-313.37. Roblamg. LX. a"d Davidson, J.D. (1914): The 45. Tsvlleneva,. V.A. (1940 The nervous system ol

anatomy of Ants; pssuleu(Oken). I8J8. Part IL. the ixodid ganglion. Zoologische JahrbucherParasitology 6: 217-256. Anatomic undt Antogenie der litre 111: 576-6202.

38. Roebdly, M.A. (121: The structure of the 46. Tsvlenseva. V.A. 119450 The nervous structure olintracellular rickeittia-like microorganisms in the ittodi synganglion (Acarina. Ixodoidea).

* ticks, and the relationsship of the micro-organisms Entomol. Rev. 44:135-142.*to the acarines and their tissues. Ph.D. Thas. 47. Witilieswomts, V.3. (1944) The nutrition 01 the*London University. central nervous system of the cockroach.

*39. Roedy. M.A. and Marzouk. A.S. (1114); The Periplaneft americans. The mobilization ultsubgienuis Persicargas (ixodoides: Argasidac: reserves. J. Esp. Dio1. 37: 500-512.Ann)s: A. (P.) arbersti central nervous system 48. Weod. O.L.; Mosisa. M.l.; Hooiptrsal., H. endanatomy and histology. J. Parasit. 76: 774-787. 3Otiffker, W. (11111): Kadam virus (Togaviridac.

40. Sait. Y. (1944):t Studies on ixodid ticks. Pan 11. Flavivlrus) infecting camel-parasitizingrhe internal anatomy in each stage of Hyalommis dronsadarilticks (Acedlitoddae)Inbeemqttsysali flava Neumann. 1897. Act Med. Saudi Arabia. J. Med. Entomol. 19: 207-208.Biol.. NiOata 9- 189-239. 49. Zalier, M.A.; Sollinan. Z.R. aid Dish. P.M.

41. Sadja. R.C.. Siarsa S.P. and Kiemn, S. (1977): An experimental study of Bormgla*(1971); Histological studies of the neural anserlum in four species of Argas ticks. 2.

stutrsI three ticks. Ityalemos wigyptium Tranistadiat survival and transovarialSharif. Rlpicephl~s sanguineus Latreille and transmission. Z. Parasitenk. 53. 213-223.'"A. 0

320

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Synganglion Histology of the Camel Tick Hyalomma dromedarii (Acari:Ixodoidea:Ixodidae).

12 PERSONAL AUTHOR(S)

Marzouk,_Aleya S.*, 'ilil, Galila M.+, Mohamed, Fatma S.A.* and Abdel-Kawy, Soht ir*i3a TYPE OF REPORT 113b TIME COVEREC, 114 DATE OF REPORT (Year, Month. a)15PG COUNT

I ROM TO -I 1966, December 116 SUPPLEI.. XRY NOTATION

Published in: Egypt. J. Histol., 9(2):309-320, 1986; ACC. No. 1499.

,7COSATI CODES 18. SUBJECT TERMS (Continue on reverse if necessary and iderify by bloc number)

FTEC GROUP SUB-GRC: Ticks; Ayaiomia dromedari'; Histology; Synganglion.

1.9 ABSTRACT (Continue on reverse if necessary and identify by block nurn.er)

The synganglion histology in female H. cicoedarii is described and compared with o. r ticks.The syngang 7ion, lying in a perigancjionic blood sinus, is formed by a fused supra--3ophagealpart-(protocerebrum, cheliceral gancdia, pedipalpal ganglia and stomodeal. pons) an( * sub-esophageal Part (4 pairs of pedal ganglia and the complex opisthosomatic ganglion). :,he syn- _N1

ganglion is enclosed within a connective tissue sheath, the neurilemma. The perineu Lumunderlies the neurilemma and is followed by the cortical region, which is separate6 Croin theneuropile by the subperineurium. Association, motor arnd neurosecretory cell types aoe the 0neurons whi-h together with 4 of 5 glial cell types, compose the cortical zone of t snganglion. Thie fifth glial cell type is only distributed in the perineurium. Five le lds offibrous horizontal connectives and vertical commissures occur in the neuropile.

*Zoolo ' Department, Faculty of Science, Ain Shamis University, Cai.ro, Egypt.+Zooloc,! Department, Faculty of science, Qatar University, Qatar.

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