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Leishmaniasis in Brazil: Xi LEISHMANIASIS IN BRAZIL: XI. OBSERVATIONS ON THE MORPHOLOGY OF LEISHMANIA OF TOE BRAZILIENSIS AND MEXICANA COMPLEXES J. J. SHAW and R. LAINSON The Wellcome Parasitology Unit, Instituto Evandro Chagas,Belém, Pará, Brazil Reprinled from lhe JOURNALOF TROPICALMEDICINE ANOHVGIENE, January 1976, Vol. 79, pp. 9-13. LEISHMANIASIS IN BRAZIL: XI. OBSERVATIONS ON THE MORPHOLOGY OF LEISHMANIA OF THE BRAZILIENSIS AND MEXICANA COMPLEXES J. J. SHAW and R. LAINSON Wellcome Parasitology Unir, IrlStituto Evandro Chagas,Belém. Pará. Brazil Introduction H6 L. mexicana amazonensis: isolated While studying strains of Leishmania isolated from a non-ulcerated lesion on the from man and animaIs in the Amazon we thigh of a man from the Paragominas noted that they relI into two natural groups. region of Pará. (2057'S -47°13'W). These we called "fast" and "slow" strains H17 L. mexicana amazonensis: isolated (Lainson and Shaw, 1970) because of their from a lesion of a woman with ex- behaviour in culture and hamsters. It must be tensive lesions on legs and back. emphasizedthat the terms " fast" and " slow " many of which were ulcerated. from do not reter to their behaviour in mano When the River Maracá region of Amapá. examining Giemsa-stainedsmears from infected (OO20'S-51 °35'W). hamsters we gradually became aware that we H9 L. braziliensis braziliensis: isolated could distinguish the "fast" and "slow" from a single ulcer infection below strains. At first we thought that it was possibly lhe knee of a man from the Itinga due to differences in the cellular picture and region of Pará. (4°36'S -47°43'W). number of parasites, but blind tests in fields Ml678 L. braziliensis braziliensis: isolated with similar numbers of parasites and cells from a small single ulcer on the enabled us to distinguish parasites of the two thigh of a man from the Serra Norte groups, which we termed the braziliensis com- region of Pará (Lainson et al., 1973). plex (slow) and the mexicana complex (fast) (Lainson and Shaw, 1972). Clearly we were Amastigotes of the different strains were recognizingbasic morphological differences but obtained from non-ulcerated cutaneous nasal we could not be certain what they were. lesions of golden hamsters (Mesocricetus To determine the morphology of the ama- auratus). stigotes more precisely we decided to apply Previous morphological studies on Sarcocystis the classicalprocedures used to study the mor- zooites (Lainson and Shaw, 1969) showed that phology of trypanosomes. It soon became the methods of fixation and collection signifi- apparent that camera lucida drawings of such cantly effect size. Preliminary observations on small organisms were not very reliable. We Leishmania amastigotes showed that direct therefore used pho~omicrographic techniques smears from lesions were not a suitable source to determine the various mensural characters of parasitesas there was considerable variation (Shaw and Lainson, 1972) of the amastigotes. in shape depending on the pre~nc.e or absence This paper reports the basic morphological of blood. In general amastigotestended to be characters of some strains of the braziliensis and mexicana complexes. torpedo shaped when associated with blood. Least variation in shapewas noted in impression smears made from a piece of tissue from the Materiais and Methods lesion whose surface had been gently blotted The following Leishmania strains were used:- on filter paper. Smearsmade in this way were PH7 L. mexicana amazonensis: isolated fixed in Bouin's fluid and stained with a modi- from a naturally infected Lutzomyia fied Giemsa technique (Lainson, 1959;Wenyon, flaviscutellata captured in the Utinga 1926). Photomicrographs of arnastigotes were forest, Belém. (Lainson and Shaw taken on Adox KB 17 or Agfa Isopan IF 35 1968). fim. film using a Zeiss WL research micro- M22 L. mexicana amazonensis: isolated scope with a Planapo 100/1'3 oil objective and from a naturally infected Oryzomys the Zeiss 11 carnera body mounted on a sup- capito captured in the Utinga forest, port armoEach fllm included a photomicrograph Belém (Lainson and Shaw, 1968). of the scale of a slide micrometer which was The H6 H9 plastjposterior end to middle óf nucleus (I1</ PN); posterior end to middle of kinetoplast/ middle of kinetoplast to anterior end (PK/KA); posterior end to middle of nucleusjmiddle or nucleus to anterior end (PN /NA). If the PK/ PN ratio is greater than one the kinetoplast is anterior to the nucleus. if it is one then they are in the same position. and if it is less than one then the kinetoplast is posterior to the nucleus. If the PKjKA is greaterthan one then the kinetoplast is in the anterior half. if it is one then it is in the middle and if less than one it is in the posterior half. The position of the nucleus as determined by the PN /NA ratio fallows the same paliem as the PK/KA ratia. Results I I A summary of the various mensural characters I micron is given in Table I. Analysis by the F-test Fig. (i). Amastigotes of L. m. amazonensisshowing mean mensural characters of strains H6 and H9 in showed that there were highiy significant dif- diagramatic formo ferences betweenthe lengths of L. m. amazon- ensis strains. ranging from 3' 18 to 3' 72 p.m printed with each ftlm and used to determine with a mean of 3' 44 p.m.The degreeof overlap lhe final magnifications of the prints. The sizes betweenthe smalIer strain M22 and lhe larger of the organisms.organelles and their positions strain H17 was 23 per cento The overlap be- were measuredfrom the photomicrographs with tween the two strains of L. b. braziliensis was a pair of sliding callipers. The anterior end was 26 per cento the mean major diameter being taken as being the end of lhe organisms in 2. 4 p.m. The overlap between the smaller which the ftagellar vacuole occurred (Figure i). L. m. amazonensisstrain M22 and the larger Dividing forms with two nuclei or kinetoplasts L. b. braziliensis strain H9 was 12 per cento were not included in the calculations of the while the overlap of the major diameter for the mean biometrical characters.Wherever possible two groups was 13 per cento The mean area extracellular parasites were measured. The of the amastigotesof strain M22 was 5.34 p.m2 meansand standard deviations were determined and H9 was 3,78 p.m2with an overlap of area for lhe different mensural characters including of 24 per cento The PK/pN fatia was greater the major diarneter of the kinetoplast. Vari- than one for the strains of L. m. amazonensis ations within and betweenthe two groups were and approximately one for the L. b. braziliensis analysed using the F-test and differences be- strains indicating that the kinetoplast is more tween means were compared using the fatia anterior to the-nucleus in the former. The over- d (Bailey. 193:,9).The percentageof overlap of lap of the distance P- K betweenstrains M22 certain characters was estimated using the and H9 was 20 per cent and betweenM22 and M1678 15.9 per cento The PKjKA fatia in- formula C= 1/2 (xl -x2). where S is the dicated that the kinetoplast is more posteriorly S placed in the two L. b. braziliensis strains and common estimate of the standard deviation of the overlap of this character between strains the two popu1ations as ca1cu1atedfor a l-testo M22 and H9 was 28.4 per centoThe PNjNA The percentageof over1apis 1- P, where P is fatia showed that the nucleus is more centralIy the integral of the normal distribution from placed in the L. b. braziliensis strains. -C to infinity. Thus if the above gives a deviate The sizesof the nuclei of alI the strains were C= 3.24, tab1esof the normal distribution give similar ranging from 1.2 X 1.01 p.m to 1.35 P=0.9994 and the over1apis 1-0.9994=0.06 Xl' 05 p.m. The kinetoplasts of the L. b. per cent [in the original examp1e(Shaw, 1969) braziliensis amastigoteswere significantly larger ..C of 23.4" shou1dhave read ..C of 3.24 "]. than those of L. m. amazonensis.An F test of Ratios for the fo110wingcharacters were also the major diameter of the kinetoplast was signi- ca1cu1ated;posterior end to midd1e of kineto- ficant for the L. m. amazonensisstrains but not 1)1::':)7~1J TABLB I. Mensural data in microns of four strains of Leishmania mexicana amazonensis and two strains of L. bra- ziliensis braziliensis isolated froro man and wiId animaIs in the state of Pará, Brazil (Data expressed as means ::!: standard deviations and range). for the two L. b. braziliensisstrains. The overlap major (L. major). Doubt as to the validity of of the major diameter of the kinetoplast be- this differentiation was expressed by some of tween strains PH7 and M1678 was nine per cento the Russian workers but Kellina (1962), in a Amastigotes of L. m. amazonensisin general detailed study of the sizesof the parasites from had larger flagellar vacuoles. The mean men- 13 human cases of cutaneous leishmaniasis, sural characters of strains H6 and H9 are concluded that there were significant differences shown in diagramatic forro in Figure (i). - in sizes of amastigotes associated with the urban and zoonotic forms of the disease.The DiscvssioD mean sizes of parasites of the urban fornl (L. tropica) were 3,33 X 1,99 p.mwhile those Such statements as "the leishmania are a of the zoonotic form (L. major) measured group of morphologically identical parasites" 4,48 X 3, 33 Jl.m. She emphasized that great (Schnur et al.. 1973) have often appeared in cale was needed in the making of the smears the literature.
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