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PUBLICATION REPORT

[APPrOved ~Pbi ~k~~10

19/88

NEUJIOSCRETORY CIT-, TYPES AND DISTRIBUiTION IN 01751) FEMALZKYALCI4MA DRONEDARI (ACARI IXODOIDEA:IXODIDAE) BYNGPINGLION

BY

A.S. MatCzouk, G.M. Khalil, and Z.E.A. Darwish

U.S. NAVAL MEDICAL RESEARCH UNIT NO. 3'CAIRO, ARAB REPUBLIC OF EGYPT)

FPO NEW YORK 09527

4393 0 7 05 3

r-r

Journal of the Egyptian Society of Parasitology. vol. 17 No. 1. 1987.

NEUROSCRITOBY CELL TYFES AND DISTRIBUTION

IN UNFED FEMALE HYAWMMA DROMEDAII

(ACARI: IXODOIDEA: IXODIDAE) SYNGANGLION*+I

By

A.S. MARZOUKz, G.M. KHALIL, and Z.E.A. DARWISH,

1. Medical Zoology Department, United States Naval MedicalResearch Unit Number Three (NAMRU-3), co AmericanEfbay, Cairo, Egypt.

2. Zoology Department, Faculty of Science, Ain Shams Univer-sity, Cairo. Consultant, Medical Zoology Department, NAM-RU-S.

3. Present eddress: Zoology Department, Faculty o1 Science,Qatar University, Doha, Qatar.

4. Zoology Department, Faculty of Science, AI-Azhar Univer.sity for Girls, Cairo. Guest Investigator, Medical ZoologyDepartment, NAMRU-,

* From Research Project 3M161102BSIO.AD.424,. Naval Me-dical Research and Development Command, Naval MedicalCommand, National Capital Region, Bethesda, Maryland.

.Supported in part by contract RRO41-26-01, NR205-040 withthe Office of Naval Research, Microbiology Program, NavaiBiology Project, Arlingto4, Virginia. The opinions andassertions contained herein are the private ones of theauthors and are not to be construed as official or as reflect-ing the views of the Department bf the Navy of' of thenaval service at large.

+ Dedicated to late Dr. Harry Hoogstraal who initiated andsupported this work before hd passed away on 24 Februliry

1986.

-41-

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ThirteertJ4SC)ypes have been observed in the unfed femaleHyeSorna dromedarii synganglion. Distribution of these typeswas based on cell shape, size, and staining reaction of theircontents. The NSC are grouped in 13 centers, namely proto-cerebral, cheliceral, stomodeal pons, palpal, oesophageal,l glo-bular, olfactory glomerular, four pedal, opisthosomak and post-oesophageal. Each center contains one or more cell types. 1'.

INTRODUCTION

Gab& (1955) was the first to describe secretory activitywithin nerve cells in the synganglion of the argasid ticks Orni-thodoros (Pavlovskyella) erratics Lucas and 0. (Alveonamua)kahorenai8 Neumann. Subsequently, neurosecretory cells (NSC)have been described in the ixodids Boophilus (Uroboophilua)microplu8 (Canestrini) (Binnington and Tatchell, 1973), Derma-centor (D.) pictus Herm (Ioffe, 1964), D. (D.) variabilis Say(Obenchain, 1974a, b; Obenchain and Oliver, 1975), Hyalomma(H.) a8iaticum Schulze and Schlottke Ixodes (I.) ricinu8 (Lin-naeus) (loffe, 1964), 1. (.) peruicatus Schulze (Panfilova,1980), Rhipicephus (R.) aanguiseua (Latreille) (Chow andWang, 1974), and the aigasids Arga, (Persicargas) persicusOken (Eisen et al., 1973), 0. (0.) savignyi (Audouin) (Evansand Solomon, 1977) and 0. (P.) parkeri Cooley (Pound andOliver, 1982).

- The first atteipt to classify the NBC into distinctive typeswas in 0. (0.) moubata Murray (Eichenbergr, 1970). Later,Obenchain (1947) presented an arbitrary classification schemefor the NSC in D. variabilis and Gabbay and Warburg (1977)in 0. (P.) tholosani.

In H. (H.) dromedarii Koch, changes in the neurosecretorysystem have been correlated briefly with moulting and oviposi-tion (Dhanda, 1967). In this study, we describe in detail theNSC types and their distribution in the synganglion of unfedfemale H. dromedarii to provide background data essential 'or

-42

subsequent studies on changes in these cells associated withfeeding and mating.

MATERIAL AND METHODS

Adult H. dromedarii were colonized in the NAMRU-3 Me-

dical Zoology laboratory from females collected from camels in

the Imbaba market, Giza, Egypt, using domestic rabbits as hosts.Methods of feeding and rearing were those of Berger et al.(1971) at 28 =- IOC and 75% relative humidity. All animal care

and manipulations were in accordance with the Animal Welfare

Amendment of 1976 (PLP14-279) with subsequent amendments.

For histological examination, ticks were dissected on the

day of moulting in a 0.77 saline solution and the synganglia

were fixed in Bouin's fluid, dehydrated and double-embedded in

.celloidin-paraplast. Five to 7 p-thick serial sections were stainedusing Mallory triple stain (MT) (Pantin, 1959), Ewen's (1962)paraldehyde-fuchsin (PF) and Gomori's (1941) chrome-haema-toxylin-phloxine (CHP).

Cellular diameter of 6-10 specimens were measured.The surface area of sections of oblong cells

axbwas calculated using the formula A = 7 x - where A is the

4area, a is the small diameter and b is the large diameter. The

mean and standard error were then calculated for each cell type

-and the data were compared using the Student's t-test.

RESULTS

Thirteen NSC types (I-XII) were observed in differentlocations in the synganglion. Distinction of these types was'based on cell shape, size and staining reaction of their contents

as illustrated in Table 1.

Type I : These NBC are the largest of all cell types (P <,0.001). The monoaxonal cell is oval, and contains several peri-

-43 -

-. , =.....=.=..... nm mm - ==ml i lll I I I

pheral vacuoles and a large central nucleus with 3 nucleoli(Fig. 1). Some of the densely-distributed neurosecretory gra-

nules (NSG) clump togethcr forming larger aggregates aroundthe nucleus and others are traced in the axon.

Type 11 : These NSC are oval with a central nucleus having2 nucleoli. Two subtypes are observed according to cell size andNSG amount and distribution. Subtype Ila cells have very finegranules enclosed in areas of clear, unstained cytoplasm whichgives the cytoplasm a vacuolated appearance (Fig. 1). Smallersubtype Ub cells (P < 0.02) contain coarse granules with largeaccumulates on one side of the nucleus (Figs. 1, 4).

Type III : These NSC are large, oval and their nuclei con-tain 2 nucleoli. Two subtypes are distinguished according to cellsize and NSG shape and distribution. In the larger subtype IIacells (P < 0.01) the cytoplasm contains several large vacuolesand the coarse NSG aggregate more densely at one side of thenucleus (Fig. 1). The subtype Ib contains clamps of flocculantgranules uniformly distributed in the cytoplasm (Figs. 1, 5).

Type IV : These are monoaxonal, more cr less oval, eachcontaining a large vacuole and an eccentric nucleus with 2nucleoli (Figs. 1, 6). The fine NSG mainly aggregate near thecell periphery and in the axon.

Type V : These are large, irregularly-shaped cells, eachcontaining 2 large vacuoles and a large central nucleus with onelarge nucleolus (Figs. 2, 7). The fine floculant NSG aggregatemostly at 2 sides of the nucleus.

Type VI : These are small, oval cells, with eccentric, sphe-rical nuclei. Three subtypes are distinguished according to NSGdistribution. and or cell size. In subtype Via, the NSG areuniform -and evenly distributed (Figs. 2, 8). In subtype VIbsome of the fine NSG accumulate around the nucleus (Fig. 2).In subtype VIc large aggregates forming droplets are scattered*among the fine granules (Fig. 2).

-44-

i!

Type VII: These are the smallest (P < 0.001) NSC. Theyare rounded with relatively large nuclei and large NSG. Twosubtypes VIla and VIIb are observed according to NSG distri-bution ; the granules are scattered evenly in the cytoplasm ofthe former and irregularly in the latter subtype (Fig. 2).

Type VIII : These are monoaxonal cells in which 2 sub-types are observed according to NSG distribution : the NSG areevenly scattered in subtype VIlla (Figs. 2. 8) and accumulatein one side of the cell in subtype VIh (Fig. 2).

Type IX : In these oval celo, 2 subtypes are distinguishedaccording to cell size and NSG distribution. The NSG are en-closed in areas of clear, unstained cytoplasm in the smaller(P < 0.01) subtype IXa cells (Figs. 2, 9) and are distributedmore or less evenly in the cytoplasm of the larger subtype IXbcells (Figs. 2, 10).

Type X : These are oval, unipolar cells with a homogeneous.cytoplasm which contains a peripherally located vacuole and acentral nucleus with 2 small nucleoli. The fine to coarse NSare irregularly distributed in the cytoplasm (Fig. 2).

Type XI : These are oval cells with relatively large, roundedcentral nuclei, each containing 1 or 2 large nucleoli (Fig. 3).The cytoplasm contains several small peripherally-locatedvacuoles. The fine NSG are dist4buted throughout the cyto-plasm with heavy aggregates at one side of the nucleus.

Type XII : These are irregularly-oval cells with oval

eccentric nuclei. Each containing 3 large nucleoli (Fig. 3). Thefine NSG are scattered throughout the cytoplasm with colloidaldroplets among them.

Type XIII : Two subtypes are observed in these irregularly-outlined cells according to their size and number of axons. The-relatively smaller (P < 0.01) polyaxonal subtype XIa cellsand the large monoaxonal subtype XHIb cells possess a centralnuclei, each with 2 nucleoli (Figs. 3, 11). The cytoplasm con--

45 -

tains several small vacuoles mainly at the cell periphery. TheNSG are more heavily aggregated around the nucleus.ABC distribution in the aynganglion centers

Within the different cortical regions of the syngangiion.the NSC are grouped in 13 centers. These groups are bilaterallysymmetrical, each containing one or more cell types. Figure 12and Table 2 illustrate the distribution of 22 neurosecretory)groups in these centers.

Protocerebral center. This center includes 4 pairs of bila-terally symmetrical groups. Group 1, is the dorsal optic which.Conssts of 2 or 3 type VIIa and 2 or 3 type X cells. Group 2, isAnteroventral and consists of 4-6 NSC. It includes 2 type Ilbxells, 1 or 2 type VIc cells and one type X cell. Group 3, is pos-terodorsal and consists of one type VIMh and 2 type XU cells.Group 4, is posterolateral and consists of 4 cells type IV, IX,X and XII and 2 type VIIb cells.

Cheliceral center. This center contains 3 pairs of bilaterallySymmetrical groups. Group 5, is anterodorsal and consists of 2type VIIa cells and one type IXa cell, Group 6, is ventral andconsists of only one type Xa cell, while Group 7, is lateral and.consists of 4 NSC of types Via, VIIa, Villa and IXb.

Stomodeal pons center. Two pairs of bilaterally symmetricalgroups occur in this center. Group 8, ip dorsal and consists of2 type Vila cells, and Group 9, is ventromedial and consists of

2 type VIa cells.

Palpal center. This center contains 2 symmetrical groups..Group 10, is dorsolateral and consists of one type Mila cell, and3 or 4 type VIIb cells, and Group 11, is ventrolateral and con-sists of 2 type V cells and one type VIIa cell.

Oesophageal center. This center includes the bilaterallysymmetrical Group 18, which lies ventrolateral to the oesophagusAnd consists of I or 2 type VIa, 2 or 3 type V11b, and 2 or 3type XI cells.

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12FIGURES

Figs. 1-3 :Diagr-ammatic illustrations of the neuroseae-.tory cell type I-XII in unfed female Hyalommadran~rf.

pipl. 4-11 : Transverse sections in the synganglion ofunfed female H!/alomma droviedaris showing differentne ry cell types (arrows). X1500.

4. Type l b. 5. Type Mb. 6. Type M. 7. Type V. 8.Types VIM (a) and Via (b). 9. Type M~. 10. Type

M~. 11. Type X/Ha.Fig. 12 : Diagrammatic illustrations (W-O showing the

distribution of the neurosecretory groups (1-22) inthe different ganglia of unfed female /Hyalmmadroneftra mynganglion. ChG, cheliceral ganglia ; E,

oeophagus; OG, opiathosomatic ganglion ; Of&Kolfactory knots ; OpG, optic ganglia ; P 1-4, first to

foth tel ia; Pp G. pia gnl pro

toewebrm ; St, stomodeal p*7m.

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. . .. torylcellltypes -llI/ in liled/feIaleI HIa I mmI

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G6bular center. This center contains one pair of bilaterallysymmetrialroup 13, which is anteroventral to the globularcells and consists of 1 type VIla and 1 or 2 type VIIb cells.

Olfactory glomeruiar center. This center contains one pairof bilaterally .symmetrical Group 14. This is ventral and.-con-sists of 2 type lib, 2 type Vllb, and 2 type Xfllb cells.

Pedal centers. In each .of the 1st, 2nd, 3rd arid 4th pedalgaeglia occur 2 pairs of bilaterally synimetrical -groups (15 and1&):with a similar pattern of NSC distribution. Group 15, liesdorsolateral in each pedal ganglion andiconsists of 5 cells, typesIMes, mb, VIb, V l and VI1b. Group 16, lies ventrolateral in

- each pedal ganglion and c6nsist9 of 2 type VIla cells and 2 typeVUb cells. -

Opisthosoma! cer.ner. This center con~airs 3 unpaired and2 paired, bilaterally symmetrical groups. The unpAired Group 17is dorsommedial and consists of 5-7 NSC, 2 type, ib, one typeIV, 1 type- V1a, 2 type VIro,' and one type Vlb .cells. Thepaired Group 18, is dorsolateral and consists of one type llbT"i'andone'type Xfa cell. Group 19, is unpaired and vettromedial;it consists of 7-9 cells, 1 or -2 type I 5 type Vhs. 'one -type XfIlcells and one -type XII~b. Group 20 is paired and ventrolateral'<it consists of 6-7 NSC ; 3 or 4 cells type VIIa and 3 type VIIIb.Group 21, is unpaired and posterior ; it consists of 3 type Ha,2"type VIla, 2 type VlM, one type XIIIa and 2 type XIIb cells.

Poetoesophageal center. This center contains a pair ofbilaterally symmetrical Group 22, which consists of 4 type lib,one type ha and 3-5 type VHa cells.

DISCUSSION

Thirteen different NSC types have been observed in theunfed female H. dromedarii synganglion according to the cellshape, size snd staining reaction of their contents.: Quantita-tively, these types are similar to those described in the ixodid.Permacextor varibilis (Obenchain, 1974b) and the argasid

C.., -56 -~....... ....... ..'.......... .. , ... :!

- ,m--.mmm ] mm mmm mm mm []mm m m.m m

-7

Ornithodore thoomi (Gabbay and Warburg, 1977). Only 2types, A and B, were found in 0. moubata (Eichenberger, 1970)'one type in A. persics (Elsen et al., 1973) and 2 types . and Bin R. suguhds (Chow and Wang, 1974).

The form, quantity andor distribution of the NSG in mostof the different NSC types in H. dromedarii differ from thoseobserved in other ixodid ticks (Obenchain, 1974b). Nevertheless,the NSG chromophilla In certain cell types in H. dromedarii issimilar to that observed in certain NSC types of D. variabilis(Obenchain. 1974b) and 0. thoioani (Gabbay and Warburg,1977), yet most NSC in H. dromedarii are larger than thosedescribed in D. variabis. Types I, 1I, VII and XI among H.dromedarii NSC contain NSG which stain purple to dark purplewith PU, and are thus, similar to types I, VII, IX and X in D.vaablZs (Obenchain, 1974b) and type II in 0. thoozani (Gab-bay and Warburg, 1977. This chromophilia indicates that thesecretory material contains high levels of cysteine (Obenchain,1974b).

Comparison between PF staining reactions of NSG in celltypes of both H. dromedarit and D. variabfis (Obenchain, 1974b)indicates that types VI in the former stains purple to reddishorange as type V in the latter, while type VII in the formerresembles types H and XI in the latter ; they all stain dark.reddish purple. Types H, IX, and XIII in H. dromedarii stainbluish purple as types IMI, IV and VI in D. variabilis and typesI and V in 0. tholoani (Gabbay and Warburg, 1977). NSGstaining reddish or bluish purple with PF seem to have a highercarbonyl content than those which stain dark purple (Obenchain,1974b).

Types IV, V, X, XII in H. dromedarii contain an acidophilicsecretory product with a staining affinity to the orange G inPF technique, thus simultating type XIIa in D. variabils andtype X in 0. tholosami (Obenchain, 1974b; Gabbay and War-burg, 1977). Acidophilia persisting after oixdation may be attri-buted to proteins rich In tryptophan (Rasbe and Monjo, 1970 ;Baudry and Baehr, 1970). Cytoplasmic affinity for the light

57 -

• -. rmm n nn

green counterstain of CHP technique does not exist in H. dro-medark, although present in D. variabdie (Obenchain, 1974b)and 0. thoiobni (Gabbay and Warburg, 1977).

Grouping of NSC varies in ixodids, being fifteen groups inB. sscrapbu (Binnington and Tatchell. 1973), eighteen groupsin D. pictus (loffe, 1964), eighteen centers in D. warisbilia(Obenchain and Oliver, 1975), and eighteen centers in I. per-sacstus (Panfilova, 1981). Dhanda (1967) recognized eighteenNSC centers in H. dromedarii synganglion, without specifyingtheir locations, but here we observed only thirteen centers con-taining 22 groups in this tick.

The fifteen and eighteen groups of NSC observed in B.micropho and D. pictua, respectively (Binnington and Tatchel,1973 ; loffe, 1964) are recognizable within the thirteen NSCcenters here described in H. dromedarii as well as the eighteencenters in D. variabilis (Obenchain and Oliver, 1975). Differen-ces in the centers number in the two latter ixodids is mainlydue to the inconsistent classification of NSC in the cheliceral,pedipalpal, and opithosomal ganglia and stomodeal pons, intomore than one center. In D. varia (Obenchain, 1974b) eachneurosecretory component contains single typed-cells whereas inH. dromedarii it contains one or more cell types. As suggested

by Obenchain (1974a) the small numbers of cells in each neuro-secretory group seems to reflect an inherent economy whichis also expressed in the condensation and miniaturization ofacarine central nervous system.

In the present study, NSC presently demonstrated in theunfed female H. dromedarii require further study to elucidatetheir activity in hormonal secretions during and after feedingand their role in the regulation and coordination in the physio-logical events occurring during these phases. Studies on theseparameters are underway.

ACKNOWLEDGEMENTS

We are greatly indebted to Professor Mohamed A. Roshdy,

-58-

Department of Zoology, Ain Shams University and ScientificConsultant at Medical Zoology Department, NAMRU-3, Cairo,for his encouragement and avluable comments, and for criticallyreading the manuscript. Thanks are also due to Dr. William H.Dees, Head, Medical Zoology Department, for his interest inthis work.

REFERENCES

Baudry, N. and Baehr, J.C. (1970) : E-ude histochimique descellules neurosecretrices de 'ensemble du system nev-veux central de Rhodnius prolixus Stal (Heteraptera,Reduviidae). C.R. Acad. Sci. Paris 270:174-177.

Berger, R.S., Dukes, J.C. and Chow, Y.S. (1971) : Demonstrationof a sex pheromone in three species of hard ticks. J.Med. Ent. 8:84-86.

Binnington, K.C. and Tatchell, R.J. (1973) : The nervous systemand neurosecretory cells of Boophilus microplus (Aca-rina : Ixodidae). Z. Wiss. Zool. 185:193-206.

Chow, Y.S. and Wang, C.H. (1974) : Neurosecretory cells andtheir ultrastructures of Rhipicephalus &anguinew. (La-treille) (Acarina : Ixodidae). Acta Arachn. 25:53-67.

Dhanda, V. (1967) : Changes in neurosecretory activity at dil-ferent stages in the adult Hyalomma dromedarii Koch1844. Nature 214:508-509.

Eichenberger, G. (1970) : The central nervous system of Ort-thodoros moubata (Murray) (Arachnida : Acari : Ixo-doidea : Argasidae) and its postembryonic development.Acta Trop. 27:15-33.

Eisen, Y., Warburg, M.R. and Galun, R. (1973) : Neurosecretoryactivity as related to feeding and oogenesis in the fowl-tick Argas persicum (Oken). Gen. and Comp. Endocrinol.21:331-340.

Evans, A.A. and Solomon, K.R. (1977) : Neurosecretion inOrnithodoros savignyi (Audouin) (Ixodoidea : Arga-sidae). The distribution of neurosecretory cells in thebrain. J. Vet. Res. 44:127-130.

Ewen, A.B. (1962): An improved aldehyde fuchsin staining

- 59 -

technique for neurosecretory products in insects. Trans.Amer. Mier. Soc. 81:94-96.

Gabbay, S. and Warburg, M.R. (IT) : Neurosecretory activityas related to feeding, mating, and oogenesis in thefemale cave tick Orrnthodoroa thokjsssw (Acari: Ar-gasidae). J. Insect Physiol. 22:1291-1301.

Gabe, M. (1055) : Histological data on neurosecretion in arach-nids. Arch. Anat. Micr. Morph. Ezxp. 44:351-383.

Gomori, G. (1941) :Observations with differential stains ofhuman islets of Langerhans. Amer. J. Path. 17:395-406.

loffe, I.D. (1964) : Distribution of neuroaerevtory cells in thecentral nervous system of Dermacestor pictus. Herm(Acarina, Chelicerata). Dohi. Akad. Nauk. S.S.S.R.,154:229-=3.

Obenchain, F.D. (1974a) : Structure and anatomical relation-ships of the synganglion in the American dog tickDermacentor vaniabilis (Acari : Ixodldae). I. Morph.142:205-223.

Obenchain, F.D. (1974b) : Neurosecretory system of the Ame-rican dog tick Dermacentor variba (Acari : Iodi-dae). 1. Diversity of cell types. J. Morph. 142:433-446.

Obenchain, F.D. and Oliver, J.H. (1975) : Neurosecretory systemof the American dog tick Dermacentor variabilis (Acari:Ixodidae). IU. Distribution of secretory cell types,axonal pathways and putative neurohemal neuroendo-crine associations ; comparative histological and ana-tcmical implications. J. Morph. 145:269-294.

Panfilova, I.M. (1980) : Changes in the neuro-endocrine kystem1) female Ixodes peraidcatus during the feeding period.

ni.Zh. 59:851-858.Pantin, C.p.A. (1959) : Notes on microscopical techniques for

---1cgists. Cambridge University Press.Pound, I.M. and Oliver, J.H. (1982) : Synganglial and neuro-

secretory morphology of female Orrnthodoros parkeri(Cooley) (Acari : Argasidae). I. Morph. 173:159-177.

Raabe, M. nd Monjo, D. (19T0). Recherches histologiques ethistochimiques stir Is neurosecretlon chez Ia Phasme.Chlumius extradenitalus. Les neurosecretions de typeC. C.R. Acad. Sci. Paris 261:4240-4243.

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. Egypt. Soc. Parasto 17 (1) PaPer No 5

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11. TITLE (Include Security Clasicatlon)Neurosecretory Cell Types and Distribution in Unfed Female Hyalomma dromedarii (Acari:Ixodoidea.Ixodidas) Syngariglion. (UNCLASSIFIED)12 PERSONAL AUTHOR(S)Harzouk , A.S.*, 10halil, r.M.* and Darwish, Z.E.A.+13a TYPE OF REPORT 13b TIME COVERED 14. DATE OF REPORT (YearbothDay) iS 5P AGE COUNT

IFROM TO _ 1987 21

16 SUPPLEMENTARY NOTATION

Published in: J. Egypt. Soc. Parasitol., 17(l):41-61, 1987; Acc. No. 1500.

17 COSATI CODES 18 SUBJECT TERMS (Continue on reverse if necessary and identify by block number)FIELD GROUP SUB-GROUP Ticks; flyalomma dromedarii; Neurosecretory cell;

I I I Distribution; Types.19. ABSTRACT (Continue on reverse if necessary and identify by block number)

Thirteen NBC types have been observed in the unfed female Ryaloa dromedarii synganglion.Distribution of these types was based on cell shape, size and staining reaction of their-contents. The NBC are grouped in 13 centers, namely protocerebral, cheliceral, stomodealpons, palpal, oesophageal, globular, olfactory glosserular, four pedal opisthosomal and

- st-oesophageal. Each center contains one or more cell types.

CZoology Department, Faculty of Science, Ain Shams University, Cairo, Egypt.*~Present address: Zoology Department, Faculty of Science, Qatar University, Qatar.

+ Zoology Department, Faculty of Science, Al-Azhar University for Girls, Cairo, Egypt.

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