FARELERDE PAPiLLA FUNGiFORMisLERiN POSTNATAL GELi~iMi UZERiNE I~IK VE TARAMALI ELEKTRON MiKROSKOBiK (SEM) incelemeler*

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1 FARELERDE PAPiLLA FUNGiFORMisLERiN POSTNATAL GELi~iMi UZERiNE I~IK VE TARAMALI ELEKTRON MiKROSKOBiK (SEM) incelemeler* letlik Merkez Veteriner Kontrol ve Ara~t1ITnaEnstitiisii Ankara - TORKiYE 2Flrat Oniversitesi Veteriner Fakiiltesi Anatomi Anabilim DalI Elazlg - TORKiYE Light and Scanning Electron Microscopic (SEM) Investigations on Postnatal Development of the Fungiform Papillae in the Mice Summary This study was carried out to examine light and scanning electron microscopic (SEM) changes in the fungiform papillae of mice during postnatal period. In this study, the tissue samples collected from the tongues of mice that were 0, 15, 30, 60, 90, 120, 150, and 180 days old during postnatal period were used to be material. Collected tissue samples were examined by light and scanning electron microscopy. Fungiform papillae were present in newborn mice but mature taste buds were seen in postnatal 2 days old animals. It was seen that the fastest increase in 0-15 days period and papillae completed their developments in 120 days old animals with the average length of IJ.m and width of IJ.m. The average length and width of the taste buds that were completely developed in 90 days old mice were IJ.m and IJ.m, respectively. In the scanning electron microscope, one taste pore was seen in the middle of the top of the papillae as of 15 th day of the age. By the examination of higher magnification micropits and microridges were seen on the epithelial surfaces of the tissues samples of all age group animals. As a result, it was found that fungiform papillae, which seen the first mature taste buds in postnatal 2 days old animals and were developed fast at the beginning (0-15 days) of the postnatal period. Ozet Bu yali~ma farelerde postnatal donemde papilla fungiformis'lerde meydana gelen,l~lk ve taramali elektron mikroskobik degi~imleri incelemek amaclyla planlanml~tu. Ara~tlrmada postnatal donemin 0, 15, 30, 60, 90, 120, 150 ve 180. gtinlerindeki fare dillerinden alman doku omekleri materyal olarak kullamldl. Yeni doganlarda papilla fungiformis'ler mevcuttu fakat porus'u te~ekkiil etmi~ tat tomurcuklanna postnatal 2 giinliiklerde rastlamldl. Papilla'lann ortalama olarak uzunlukta IJ.m, geni~likte de IJ.m ile geli~imini 120 giinliiklerde tamamladlklan ve en hlzli artl~ orammn ilk 15 giinliik donemde oldugu goriildii. Tat tomurcuklanmn ise 90 giinliiklerde geli~imlerini tamamladlklan tespit edildi ve snaslyla uzunluklan IJ.m, geni~likleri ise IJ.m olarak belirlendi. Taramah elektron mikroskopta, 15 giinliiklerden itibaren papilla'lann tepesinin tam ortasmda bir adet tat porus'u gozlendi. Papilla fungiformis'ler yuzeyinden yapi1an ytiksek btiytiltmelerde, btittin gruplarda ytizey epitelinde yok ktiylik yukur (Micropit) ve yok kiiyiik yoontilara (Microridge) rastlamldl. Sonuy olarak, papilla fungiformis'lerdeki olyiimlerde en hlzli artl~m ilk 15 giinliik donemde oldugu, bu donemden sonraki artl~m 120. giine kadar yava~ bir hlzla devam ettigi ve porus'u te~ekktil etrni~ fonksiyonel ilk tat tomurcuguna postnatal 2. giinde rastlandlgl tespit edilmi~tir. Anahtar Kelimeler: Fare, papilla fungifonnis, geli~im, taramah elektron mikroskop (SEM)

2 Giri~ Memelilerde papilla fungiformis'ler, dilin tist yiizunde ve yogunlukla Uy klsmmda, papilla filiformis'ler arasmda yer ahrlar (5,10,22-24). Mouse ve hamster gibi kemiricilerde papilla fungiformis'ler, dilin ust yiizunun ortasmdaki oluga paralel ve iki tarafh simetrik malar ~eklinde bulunurlar. Fakat bu olukta papilla'lar yer almazlar (24, 28). Papilla fungiformis'ler, prenatal donemde mouse'da ilk defa 13. gtin (28) veya gtinde (15), ratlarda 15. gtin (7) veya 21. gtinde (5), hamsterde ise 12. gunde (32) ~ekillenir. Papilla fungiformis'lerin innervasyonu, V. c;;ift beyin siniri olan n. trigeminus' un kolu n. lingualis ve VII. c;;iftbeyin siniri olan n. facialis'in duyusal kolu chorda tympani tarafmdan saglamr (7). Rat, mouse ve hamsterde papillanm epitel tabakasmm en tepesinde bir adet tat tomurcugu bulunur ve bir.tat porus'u ile aglz bo~luguna ayillf (5,16,23,25,31). Bununla birlikte insan ve maymun gibi primatlarda bir papilla fungiformis'de ortalama tat tomurcugu (3,4,33) ve kedide 7-17 tat tomurcugu (29) yer ahr. Rodentia'dan, oklu kirpide (19) ve keseli slyanda (18) ise papilla fungiformis'ler tat tomurcugu iyermezler. Fare ve ratlarda papilla fungiformis'lerin prenatal ve erken postnatal donemde geli~imi ile ilgili taramah elektron mikroskobik yah~malarda yiiksek btiytiltmelerde epitel hticrelerin belirgin oldugu ve bu hucreler yiizeyinde yok kuyuk ylkmtl ve yukurlarm bulundugu gozlenmi~tir (11,13,14). <;ah~mada, postnatal donemde fare papilla fungiformis ve tat tomurcuklanndaki l~lk ve taramah elektron mikroskobik degi~imlerin incelenmesi amaylanml~tlr. Materyal ve Metot <;ah~mada, dogum gunu "0" kabul edilmek uzere 0, 15, 30, 60, 90, 120, 150 ve 180 gtinltik donemler halinde top lam sekiz grup olu~turuldu ve her grupta 8'er fare kullamldl. Aynca papilla'daki tat porus'lanmn ayllma zamamm tespit etmek ic;;in0-10. gunler araslllda her gun iyin 2 adet olmak uzere top lam 84 fare kullamldl. Hayvanlar ether ile anesteziye ahndl ve olduriildu. Alman doku omekleri %lo'luk formaldehit solusyonunda tespit edildi. Elde edilen parafin bloklardan 5-7 mikrometre kalmhgmda kesitler her ya~ grubunda dilin dorsal'inden ventral'ine transversal olarak ahndl. Bu Kesitlere Mayer'in hematoksilen-eosin (21) boya yontemi uygulandl. Mikrometrik olyumler okuler mikrometre yardlmlyla ahndl. Resimler Nikon marka ara~tlrma mikroskobunda c;;ekildi. Terminolojik ifadelerin yazlmlllda Nomina anatomica veterinaria (26) ve Nomina histologica veterinaria (27)' dan yararlamldl. Taramah elektron mikroskobik inceleme iyin her gruptan iki farenin dili kullamldl. Bunlar bir buron halinde serum fizyolojikte ylkandl. Hazlrlanan doku omekleri fosfat tamponlu %3'ltik gluteraldehitte (ph:7,3) ilk tespit i~lemine tabi tutuldu. Daha soma aym tamponda ylkanan dil omekleri %1 'lik ozmiyum solusyonunda 37 C' de 1,5 saat stireyle son tespit yapiidl. Bu i~lemden soma dil yiizeyindeki ekstraselluler mukus tabakasml uzakla~tlrmak amaclyla 3 N'lik HCI'de 60 C slcakhkta 20 dakika bekletildi (13, 14). Elde edilen dil omekleri alkol ve amil asetat serilerinden gec;;irilerek 31 C slcakhk 70 bar basmyta 1,5-2 saat stireyle kritik nokta kurutucusunda (critical point dryer) kurutuldu. Kurutulan diller stub lara yapl~tlf1ldl. inceleme oncesinde, stublardaki dil omekleri VG Microtech Polaron SC500 sputter coater cihazlllda 0.05 Torr basmy altmda altm kaplama yapildl. Hazlrlanan numuneler Jeol JSM 5600 marka taramah elektron mikroskopta 5-15 kv arahgmda incelenerek fotograflar alllldl ve degerlendirildi. Bulgular Papilla fungiformis' lerin, dilin list ytiztinde man tar biyiminde ylkllltilar olarak papilla filiformis'ler araslllda daglldlklan goriildu. Dilin ucundan, ogurucu di~ler arasmdaki ylkmhsma kadar olan bolgede yer alan bu papilla'lar, yo gun olarak dilin Uy klsmlllda ve kenarlannda bulunmaktaydl. Sulcus medianus'a paralel simetrik malar ~eklinde bulunan papilla fungiformis'lere bu olukta rastlamlmadl. I~lk mikroskobik bulgular: Papilla fungiformis'ler yok kath yassl bir epitel ile ortuluydu. Papilla'lann tepesindeki epitel. kallllhgl, yanlanndaki ve papilla'lar arasllldaki epitel kallllhgmdan daha inceydi. Yeni doganlarda papilla'mn tepesinin tam ortasmda bir adet tat tomurcugunun (Caliculus gustatorius) belirginle~meye ba~ladlgl ancak bunun tat porus 'una (Porus gustatorius) sahip olmadlgl goriildu (~ekil 1). Papilla'larda porus'u ~ekillenmi~ (olgunla~ml~) tat tomurcuklanna postnatal 2 gunluklerde rastlamldl (~ekil 3). I~lk mikroskobunda tat tomurcuklannda aylk (gustatorik) ve koyu (destek) boyah'iki tip tat tomurcugu hucresi goriildu (~ekil 4). Papilla uzunlugu ve geni~ligi yeni doganlarda slraslyla fj.m ve fj.m iken 15 gtinltiklerde hlzh bir artl~la uzunlukta /lm, geni~likte de

3 66.67 /lm'ye ula~ml~tl. Papilla'nm bir onceki doneme gore uzunlukta %121.25, geni~likte de % arttlgl goriildii ve bu en hlzh artl~ oram olarak dikkati <;:ekti.daha sonraki donemlerde papilla'nm uzunluk ve geni~liginde kiiyiik artl~lann oldugu goriildii ($ekil 2). Papilla'lann postnatal 120 gilnliiklerde ortalama olarak uzunlukta /lm, geni~likte de /lm'ye ula~arak geli~imlerini tamamladlklan tespit edildi (Tablo 1). Sekil giinliiklerin papilla fungiformis'i, yok kath yassl epitel (e), tat tomureugu (t), tat porus'u (kahn ok), aylk (g) ve koyu (inee ok) boyaii hiiereler, epitel yiizeyde zaylf bir keratinizasyon (k). H.E. X 100. Sekil I. Yeni doganlann papilla fungiformis'i ve tepesinde belirginle$meye ba$laml$ bir adet tat tomureugu (t), yok kath yassl epitel (ok), bag doku (b). H.E. X 100. ;::= 140.9! 120 : : :.~100 ~ 80 -~~ ~ 60 E e 40 ~ 20 o o Hayvamn ya~ I (gun) --+- Pap. Uz. -Ill- Pap. Gen. Sekil 2. Papilla fungifonnis uzunluk ve geni~liginin ya~a bagh olarak artl~l. Yeni doganlarda tam olarak olgunla~ml~ tat tomurcugu bulunmadlgmdan olyiim yapllamadl, ancak 15 giinliiklerde tat tomurcugunun ortalama uzunlugu ve geni~ligi maslyla /lm ve /lm olarak tespit edildi. 90 gilnliiklere kadar artl~ gosteren tat tomurcuklanmn bu donemdeki ortalama uzunlugu /lm, geni~ligi ise /lm olarak saptandl ve tat tomurcuklannm geli~imlerini bu di:inemde tamamladlklan gi:iriildii (Tablo 1). Tat tomurcuklannm uzunluk/geni~lik oranlannm arasmda degi~tigi belirlendi. Taramah elektron mikroskobik bulgular: Taramah elektron mikroskobik incelemede yeni doganlann dilinde sulcus medianus belirgindi ve papilla fungiformis' lerin bu sulcus' a paralel simetrik slralar halinde papilla filiformis'ler arasmda yer aldlklan gi:iriildii ($ekil 5). Papilla.'lar yiizeyinden yapllan yiiksek biiyiiltmelerde tat porus'lanna rastlamlmadl. Sekil 3. iki giinliiklerin papilla fungifonnis'inde ilk tat porus'larmm goriilmesi, tat tomureugu (t) ve tat porus'u (ok). H.E. X 200. Sekil 5. Yeni doganlann dilinden gene I bir goniiniim. Dil ueu (a), Median sulcus (m), Papilla fungiformis (Pfu), Papilla vallata (Pva), intennolar ylkmti (i), Papilla filiformis (Pfi), dil k6kii (r).

4 Papilla uzunlugu (Ilm) Papilla geni~ligi(ilm) Tomurcuk uzunlugu (Ilm) Tomurcuk geni~ligi(ilm) H a y v a n 1 n y a ~ I g ii n ) tO.75 8S.2St t t t2.IS t3AS t AOt t t St t t t St S0t tl tO t tO S7tl.OS 38.69t l9t tl.09 29AOt S3t tO S0tO S6t A9t giinliiklerden itibaren papilla'lann tepesinin tam ortasmda bir adet tat porus'u gozlendi. Papilla'lar yiizeyinden yapllan yiiksek biiyiiltmelerde, biiti.in gruplarda epitel hiicrelerin slmrlan, c;ok kiic;iik c;ukur (micropit) ve c;ok kiic;iik C;lkmtI1ar(microridge) gozlendi ($ekil 6,7,8). ~ekil giinliiklerin papilla fungiformis'i. Papilla fungiformis (Pfu), Papilla filiformis (Pfi) ve tat porusu (ok). TartI~ma ~ekil giinliiklerde dil ucundan bir goriiniim. Papilla fungiformis (Pfu), Median sulcus (m), Papilla filiformis (Pfi), tat porus' Ian (ki.iyiik oklar). ~ekil giinliiklerin papilla fungiformis'inden yiiksek bir btiytiltme. Microridge (mr), Micropit (mp) ve tat porus'u (tp). Hamster (23,24), mouse (11,30), maymun ve insan (2), sincap maymunu (12) ve sincapta (6), papilla fungiformis'lerin dilin tist ytizeyinde papilla filifonnis'ler arasmda dagllan mantar bic;iminde C;lkmtIlar olduklan, hamster (24) ile mouse'da (28) dil ucundan oglitiicli di~ler arasma kadar olan bolgede yer aldlklan, yogun olarak dilin ucunda ve yan kenarlannda bulunduklan (2,6,24), papillalann dilin list yiizeyinin ortasmda bulunan oluga paralel olarak yer aldlklan ve bu olukta bulunmadlklan (24,28) bildirilmektedir. <;::ah~madada aym sonuc;lar elde edilmi~tir. Hamster (23,24,31), rat (5) ve mouse'da (25) papilla fungiformis'lerin tepesinin tam ortasmda papilla'nm uzunlugu eksenine paralel bir adet tat tomurcugu bulundugu ve bunlann bir tat porus'uyla aglz bo~luguna ac;lldlgl, oklu kirpi (19) ve keseli slc;anda (18) ise papilla fungiformis' lerin tat tomurcugu ic;ermedigi bildirilmi~tir. <;::ah~ma bulgulan okju kirpi (19) ve keseli Slc;an (18) ic;in yapllan bildirimlerle uyumsuz, digerleri (5,23,24,25,31) ile ise uyumludur.

5 Harada ve ark. (8), yeni dogan ratlarda papilla fungiformis'lerde tat tomurcuklanmn mevcut oldugunu ve bunlardan %14'iinde bir tat porus'u bulundugunu, dogumdan bir hafta soma ise tat tomurcuklannm %80'inin tat porus'lanna sahip oldugunu belirtirken, Diny ve ark. (5), postnatal 3 giinliik ratlarda tat tomurcuklannm yer aldlgml fakat tamamen geli~mi~ tat tomurcuklanna postnatal 5 giinliiklerde rastlamldlgml, Whitehead ve Kachele (32)' de prenatal dtinemde hamster papilla fungiformis tat tomurcugu ve porus'lannm geli~tigini ve postnatal birinci giinde tat porus'larmm eri~kinlerdeki degere ula~tigml, Iwasaki ve ark ise mouse (13) ve Rat (14)'da tat porusunun postnatal 7. giinde gi:iriildiigiinii bildirmi~lerdir. Ara~tIrmada yeni dogan farelerde tat tomurcugu bulunmu~ fakat porus'u te~ekkiil etmi~ tat tomurcuklanna postnatal 2 giinliiklerde rastlanml~tir. Papilla fungiformis'lerin taramah elektron mikroskobik yaplslyla ilgili olarak bu papilla'iann papilla filiformis'ler arasmda dagildlklan (1,6,10,11,17) ve uylanmn papilla filifolmis uylannm a~aglsmda yer aldlklan (17) bildirimleri, yah~mada da tespit edilmi~tir. Papilla fungiformis tat porus'lannm yiizey yaplslyla ilgili olarak Arvidson ve ark. (2) insan ve maymunda biiyiik delikli porus'larda mikrovillus gi:iriildligiinii ve bu mikrovillus'lar arasmda yomak benzeri hiicresel uzantilann yer aldlgml, Iida ve ark. (9) ise mouse, rat ve hamsterde papilla fungiformis tat porus'lannda mikrovillus ve yomak benzeri ylkmtilann gtiriilmedigini bildirmektedirler. <;ah~mada da tat porus'lannda Iida ve ark. (9)'nm bildirimine uygun ~ekilde mikrovillus ve yomak benzeri ylkmtilar gi:izlenmemi~tir. Bazl bildirimler (11,13,14,20), papilla fungiformis'ler yiizeyinden yapllan yiiksek biiyiiltmelerde epitel hiicrelerin slmrlanmn belirgin oldugunu, Iwasaki ve ark. (12) ise papilla fungiformis'ler yiizeyinde epitel hiicrelerin slmrlannm gi:iriilmedigini bildirmektedirler. Ara~tIrma sonuylan Iwasaki ve ark. (l2)'nm bildirimi ile uyumsuz, diger (11,13,14,20) bildirimlerle uyumludur. Papilla fungiformis' lerdeki epitel hiicrelerden yapilan yiiksek biiyiiltmelerde yok kiiyiik ylkmtl (microridge) (9-12,14,20) ve yok kiiyiik yukurlann (micropit) (11,13,14) yer aldlgl ~eklindeki bildirimler yah~mada da tespit edilmi~tir. Sonur; olarak, papilla fungiformis'lerdeki i:ilyiimlerde en hlzh artl~m ilk 15 giinluk di:inemde oldugu, bu di:inemden sonraki artl~m 120. giine kadar yava~ bir hlzla devam ettigi ve porus'u te~ekkiil etmi~ fonksiyonel ilk tat tomurcuguna postnatal 2. giinde rastlandlgl tespit edilmi~tir. <;ah~manm alamndaki bilgi birikimine katklda bulunacagl kanaatine vanlml~tir. 1. Agungpriyono S, Yamado J, Kitamura N, Nisa C, Sigit K, Yamamoto Y. Morphology of the dorsal lingual papillae in the lesser mouse deer, Tragulus javanicus. J Anat 1995; 187: Arvidson K. Scanning electron microscopy of fungiform papillae on the tongue of man and monkey. Acta otolaryngol 1976; 81: Arvidson K, Friberg U. Human taste: response and taste bud number in fungiform papillae. Science 1980; 209: Arvidson K, Cottler-Fox M, Friberg V. Taste buds of the fungiform papillae in Cignomolgus monkey. J Anat 1981; 133: Diny G, Girgin A, Yilmaz S. Ratlarda papilla fungiformis'in prenatal ve postnatal geli~imi. F.D. Saghk Bil. Derg 1995; 9: Emura S, Tamada A, Hoyokowa D, Chen H, Jamali M, Taguchi H, Shoumura S. SEM Study on the dorsal lingual surface of the Flying Squirrel. (Petauirsta leucogenys). Ann Anat 1999; 181: Farbman AI, Mbiene JP. Early development and innervation of taste bud-bearing papillae on the rat tongue. J Camp Neuro11991; 304: Harada S, Yamaguchi K, Kanemaru N, Kasahara Y. Maturation of taste buds on the soft palate of the postnatal rat. Physiol Behav 2000; 68: Iida M, Yoshioka I, Muto H. Taste bud papillae on the retromolar mucosa of the rat, mouse and golden hamster. Acta Anat 1983; 117: Iwasaki S, Miyata K, Kobayashi K. Comparative studies of the dorsal surface of the tongue in three mammalian species by scanning electron microscopy. Acta Anat 1987; 128: Iwasaki S, Miyata K, Kobayashi K. The surface structure of the dorsal epithelium of tongue in the mouse. Acta Anat Nipp 1987; 62: Iwasaki S, Miyata K, Kobayashi K. Scanning electron microscopic study of the dorsal lingual surface of the squirrel monkey. Acta Anat 1988; 132:

6 13. Iwasaki S, Yoshizawa H, Kawahara 1. Study by scanning electron microscopy of the morphogenesis of three types of lingual papilla in the mouse. Acta Anat 1996;!57: Iwasaki S, Yoshizawa H, Kawahara I. Study by scanning electron microscopy of the morphogenesis of three types of lingual papilla in the rat. Anat Rec 1997; 247: Kaufman MH. The Atlas of Mouse Development, San Diego: Academic Press. San Diego. 1992; Kinnamon CJ, Henzler DM, Royer SM. HVEM ultrastructural analysis of mouse fungiform taste buds, cell types and associated synapses. Microsc Res Tech 1993; 26: Kobayashi K. Three-dimensional architecture of connective tissue core of the lingual papillae in the guinea pig. Anat Embryol 1990; 182: Krause WJ, Cutts JH. Morphological observations on the papillae of the opossum tongue. Acta Anat 1982; 113: Kubota. K, Fukuda N, Asakura S. Comparative anatomical and neurohistological observations on the tongue of the porcupine (Hystrix cristata). Anat Rec 1966; 155: Kullaa-Mikkonen A, Sorvori TE. A scanning electron microscopic study of the dorsal surface of the human tongue. Acta Anat 1985; 123: Luna LG. Manuel of Histologic Staining Methods of the Armed Forces Institute of Pathology. Third Ed. Mc. Graw- Hill Book Company. Toronto, London, Miller n, Preslar AJ. Spatial distribution of rat fungiform papillae. Anat Rec 1974; 181: Miller RL, Chaudry AP. Comparative ultrastructural of vallata, foliate and fungiform taste buds of golden Syrian hamster. Acta Anat 1976; 95: Misu etta CM, Goosens KTA, Forinos I, Reacardt LF. Alterions in size, number, and morphology of gustatory papillae and taste buds in BDNF null mutant mice demon strata neurol dependence of developing taste organs. J Comp Neurol 1999; 409: Nomina Anatomica Veterinaria. Fourth ed. Prepared by the International Commitee on Veterinary Gross Anatomical Nomenclature and Authorized by the eighteenth general Assembly of the World Association of Veterinary Anatomists. Gent Nomina Histologica Veterinaria. Revised by the International Comrnitee on Veterinary Gross Anatomical Nomenclature and Authorized by the eighteenth general Assembly of the World Association of Veterinary Anatomists. Gent Paulson RE, Hayes TG, Sucheston ME. Scanning electron microscope study of tongue development in the CD-1 mouse fetus. J Cran Gen Dev BioI 1985; 5: Robinson PP, Winkles PA. Quantitative study of fungiform papillae and taste buds on the cat's tongue. Anat Rec 1990; 225: Utiyama C, Watanabe 1, Konig B, Koga LY, Semprini M, Tedesco RC. Scanning electron microscopic study of the dorsal surface of the tongue of calomys callosus mouse. Ann Anat 1995; 177: Whitehead MC, Beeman CS, Kinsella BA. Distribution of taste and general sensory nerve endings in fungiform papillae of the hamster. Am J Anat 1985; 173: Whitehead MC, Kachele DL. Development of fungiform papillae, taste buds and their innervation in the hamster. J Comp Neuro11994; 340: Zahm DS, Munger BL. The innervation of the primate fungiform papilla development, distribution and changes following selective ablation. Brain Res Rev 1985; 9: Miller n, Smith DV. Quantitative taste bud distribution in the hamster. Physiol Behav 1984; 32:

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