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Ommatidia of Blow Fly, House Fly, and Flesh Fly: Implication of Their Vision Efficiency

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Ommatidia of Blow Fly, House Fly, and Flesh Fly: Implication of Their Vision Efficiency Parasitol Res (2008) 103:123–131 DOI 10.1007/s00436-008-0939-y ORIGINAL PAPER Ommatidia of blow fly, house fly, and flesh fly: implication of their vision efficiency Kabkaew L. Sukontason & Tarinee Chaiwong & Somsak Piangjai & Sorawit Upakut & Kittikhun Moophayak & Kom Sukontason Received: 20 January 2008 /Accepted: 20 February 2008 /Published online: 15 March 2008 # Springer-Verlag 2008 Abstract This work aims to elucidate the number of ommatidia were detected in females. Dense postulate ommatidia or facets (the outwardly visible units of each appearance was detected in the external surface of the ommatidium) for compound eyes in blow flies [Chrysomya corneal lens of the ommatidia of C. megacephala, C. megacephala (F.), Chrysomya rufifacies (Macquart), Chrys- rufifacies, and C. nigripes, while a mix of dense postulate omya nigripes (Aubertin), Lucilia cuprina (Wiedemann)], appearance and variable groove array length was detected house flies (Musca domestica L.), and flesh flies (Liosar- in L. cuprina and M. domestica. The probable functions of cophaga dux Thomson) by manual counts of the corneal ommatidia are discussed with reference to other literature. spreads. The head of the fly in each species was soaked in 20% potassium hydroxide solution at room temperature for 7 days, and the clear compound eye was dissected into six Introduction small parts, each of which was placed onto a slide and flattened using a coverslip. Images of each part were Blow flies (Family Calliphoridae), house flies (Family obtained using a microscope connected to a computer. The Muscidae), and flesh flies (Family Sarcophagidae) are printed images of each part were magnified, and the total generally known to have important roles in medicine. Not number of ommatidia per eye was manually counted. For only do adults carry numerous pathogens (Greenberg 1971, males, the mean number of ommatidia was statistically 1973; Bidawid et al. 1978; Tan et al. 1997; Sukontason et different among all flies examined: L. dux (6,032) > C. al. 2007b), but larvae cause myiasis in humans and animals rufifacies (5,356) > C. nigripes (4,798) > C. megacephala (Zumpt 1965). In addition, flies found in human remains (4,376) > L. cuprina (3,665) > M. domestica (3,484). provide entomological evidence used in forensic investiga- Likewise, the mean number of facets in females was tions (Smith 1986; Goff 2000; Byrd and Castner 2001; statistically different: L. dux (6,086) > C. megacephala Greenberg and Kunich 2002; Sukontason et al. 2007a). (5,641) > C. rufifacies (5,208) > C. nigripes (4,774) > L. Ecologically, these flies inhabit not only human habitations cuprina (3,608) > M. domestica (3433). Scanning electron but also peridomestic or forested environments. Many of microscopy analysis of adult flies revealed the sexual them adapt well in several habitats ranging from urban, dimorphism in the compound eye. Male C. megacephala suburban, and rural areas to forested loci in various high had large ommatidia in the upper two thirds part and small altitudes. Examples of this are provided for the blow fly ommatidia in the lower one third part, whereas only small [e.g., Chrysomya megacephala (F.), Chrysomya rufifacies (Macquart), Lucilia cuprina (Wiedemann)] in Thailand by This work has been presented at the 21st Pacific Science Congress Sucharit et al. (1976), Tumrasvin et al. (1978), and held in Okinawa, Japan, in 12–18 June, 2007. Sukontason et al. (2003). In Japan, such information has * : : : : K. L. Sukontason: ( ) T. Chaiwong S. Piangjai S. Upakut been documented by Shinonaga (1965) for the blow fly K. Moophayak K. Sukontason [Chrysomya pinguis (Walker)] and the flesh fly Department of Parasitology, Faculty of Medicine, (Parasarcophaga albiceps Meigen). Chiang Mai University, Chiang Mai 50200, Thailand Biological information pertaining to various aspects of e-mail: [email protected] medically important flies is essential to devising control 124 Parasitol Res (2008) 103:123–131 Fig. 1 SEM micrographs of the heads of male flies showing the prominent compound eye locat- ed laterally. a C. megacephala with closed compound eyes in the dorsocentral area and having large ommatidia in the upper two thirds and small ommatidia in the lower one third. b C. rufifacies with closed compound eyes in the dorsocentral area. c C. nigripes with slight separa- tion of the compound eye in the mid central region. d L. cuprina with slight separation of the compound eye in the mid central region. e M. domestica with narrow separation of the com- pound eye in the mid central region strategies to cope with them. Adult flies can efficiently The aim of the current study was to determine the detect food using visual and olfactory cues (Anderson number of ommatidia of adult blow flies, house flies, and 2001). In vision, the compound eye is the most important flesh flies, which are flies of medical importance in organ made up of compact individual eye elements known Thailand. Moreover, the scanning electron microscopic as ommatidia. Each ommatidium is externally visible as a investigation was also undertaken to examine the external facet. The number of ommatidia varies in arthropods, such features of corneas of adults to enhance their bionomic as one in worker ants, Ponera punctatissima Roger information, which is of medical and/or epidemiological (Hymenoptera: Formicidae; Richards and Davies 1977), importance. seven ommatidia in booklice Liposcelis bostrychophilus Badonnel (Psocoptera: Liposcelidae), or 26 in booklice Psyllipsocus ramburi Sélys-Longchamps (Psocoptera: Psy- Materials and methods llipsocidae; Meyer-Rochow and Mishra 2007). A high number of ommatidia, such as ≈12,000–17,000 in some Flies Lepidoptera or 10,000–28,000 in some Odonata, has been estimated in some groups of insects (Richards and Davies Adult flies used in this study were obtained from laboratory 1977), thus enabling the high efficiency of vision (Blaney colonies maintained at the Department of Parasitology, 1977). Faculty of Medicine, Chiang Mai University, northern Parasitol Res (2008) 103:123–131 125 Fig. 2 SEM micrographs of the heads of female flies showing the prominent compound eye located laterally. Dichoptic—the separation of the compound eye in the dorsocentral area—is revealed in C. megacephala (a), C. rufifacies (b), C. nigripes (c), L. cuprina (d), and M. domestica (e) Thailand. Four species of blow fly from the family parts on a microscopic slide using a fine needle under the Calliphoridae (C. megacephala, C. rufifacies, Chrysomya compound microscope. A glass coverslip was placed onto nigripes Aubertin, and L. cuprina), house flies of the family each slide, thereby flattening each part of the compound Muscidae (Musca domestica L.), and flesh flies of the eye on a microscopic slide. Images of each part were family Sarcophagidae (Liosarcophaga dux Thomson) were obtained using a compound microscope connected to a used. Flies were reared in the manner described by computer. The printed images of each part dissected were Sukontason et al. (2004). magnified, and the total number of ommatidia per eye was manually counted. The data were compared using one-way Determination of the number of ommatidia analyses of variance, the least significant difference test (LSD), and paired t test. About 35 flies of each sex of each species were collected from the fly colonies. The head of each fly was dissected Scanning electron microscopic observation from the body using a sharp blade under the compound microscope. To soften the tissue for the dissection of the The external features of the corneas of adult flies (C. compound eye, the head of the fly was soaked in 20% megacephala, C. rufifacies, C. nigripes, L. cuprina, and M. potassium hydroxide at room temperature for 7 days. The domestica) were investigated in this study using scanning clear compound eye was then easily dissected into six small electron microscopy (SEM). For the SEM procedure, five 126 Parasitol Res (2008) 103:123–131 Table 1 Number of ommatidia of flies per site Fly speciesa Mean±SD ommatidia of male (range) Mean±SD ommatidia of female (range) Left eye Right eye Left eye Right eye Liosarcophaga dux 6,032±385 (5,373–6,857) 6,032±408 (5,211–6,750) 6,073±207 (5,737–6,555) 6,100±220 (5,702–6,678) Chrysomya megacephala 4,371±344 (3,437–4,927) 4,382±345 (3,502–4,925) 5,635±514 (4,777–6,649) 5,647±507 (4,874–6,678) Chrysomya rufifacies 5,367±385 (4,781–6,543) 5,346±387 (4,665–6,634) 5,207±244 (4,551–5,548) 5,209±235 (4,635–5,561) Chrysomya nigripes 4,801±323 (4,191–5,355) 4,795±327 (4,110–5,443) 4,778±277 (4,106–5,190) 4,771±278 (4,167–5,278) Lucilia cuprina 3,678±155 (3,343–3,888) 3,651±155 (3,259–3,899) 3,620±209 (3,306–3,907) 3,594±232 (3,208–4,100) Musca domestica 3,488±110 (3,223–3,841) 3,480±112 (3,254–3,781) 3,427±231 (2,781–3,787) 3,440±218 (2,846–3,777) SD standard deviation a n=30 for each species to ten live flies of each species were killed by placing them of numerous compact ommatidia, ranging from ≈3,400 to in a freezer for 20 min. The head of each fly was cut using a 6,100; with the manual counts shown in Table 1. sharp blade under a dissecting microscope (Olympus, No statistical differences between the number of omma- Japan). The specimens were attached by double-stick tape tidia in the right and left eyes of the same species were on aluminum stubs and coated with gold in a sputter- detected (P>0.05; paired t test). For males, the mean coating apparatus for 2 min so they could be viewed under number of ommatidia was statistically different among all a JEOL JSM-5910 scanning electron microscope (Tokyo, flies examined: L.
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