The appearance of LD1 minichromosomes of Leishmania and drug resistance

M. Navarro, G. Ortiz and M. Segovia
Servicio de Microbiología, Hospital General Universitario, Murcia.

The protozoan parasite Leishmania is the causative agent of visceral/cutaneous leishmaniosis, an important public health problem in the Mediterranean countries, Asia, Africa and Latin America (1). Moreover Leishmania has emerged as an opportunistic pathogen of HIV-infected humans. Since vaccines are not yet in sight, the main line of defense now available against Leishmania is chemotherapy. However, this parasite is able to acquire drug resistance. Although different mechanisms of drug resistance have been found (2), DNA amplification is the most important of them (3). Thus almost all the amplification events described so far in Leishmania are accompanied by drug resistance (4). By these means Leishmania obtains several copies of one or more genes which contribute to the resistance to methotrexate, arsenite, tunicamycin, vinblastine, etc. (3).
In our laboratory, differently cloned lines from the same stock of L. major have been obtained (5). Some of them showed minichromosomes of different sizes which were absent in the original stock. These linear DNAs have been shown to be DNA amplifications of the LD1 region situated in the 1.9 megabase chromosome (6).
To test if these amplifications were implicated in drug resistance, as most amplifications are in Leishmania, lines either with or without minichromosomes were grown with different drugs which usually kill Leishmania. Thus promastigotes of a cloned line either containing the 230 kb minichromosome (CO1 M+) or having no minichromosome (CO1 M_), were grown with: arsenite, methotrexate, tunicamycin or in the absence of any drug. At higher concentrations of the antibiotics (more than 100 mM) none of the lines was able to grow. At lower concentrations (1 µM), the growth of the CO1 M_ line (M_ in Table 1) was similar to that of CO1 M+ (M+ in Table 1). Hence no differences between the lines with the minichromosome and those without it were found. In addition, the minichromosomes could not be forced to appear in promastigotes lacking them by stepwise increasing of methotrexate, arsenite or tunicamycin (data not shown).
The apparent spontaneous appearance of these elements in different lines of Leishmania led us to analyze the dy- namics of the minichromosomes. For this purpose two tubes previously cryopreserved from the CO1 M_ cloned line were thawed, grown in RPMI medium and analyzed by Pulsed-Field Gel Electrophoresis (5). While one of the cultures had no minichromosome as the parental line did, the other one showed a minichromosome of 210 kb only forty days after thawing. These results reveal that the minichromosomes develop in a very short period of time: less than forty days. No previous DNA amplification in Leishmania could emerge in such a short period of time. For instance, amplification in drug resistance can be forced to occur by a stepwise selection process of one year, using increasing drug concentrations (7).

Table 1. Promastigote cultures with different drugs.

Mini

Day 1

Day 2

Day 3

Day 4

Day 7

Control

M_

75*

90

90

125

130

M+

50

75

75

80

90

Arsenite

M_

100

65

75

70

65

M+

70

75

90

100

55

Methotrexate

M_

90

70

75

60

65

M+

75

70

85

55

50

Tunicamycin

M_

75

100

110

100

110

M+

80

75

100

105

85

*Number indicate 105 cells/ml.

We can then conclude that the LD1 minichromosomes of Leishmania appear in very short periods of time and they are not implicated in resistance to drugs such as methotrexate, arsenite and tunicamycin.

ACKNOWLEDGEMENTS
Part of this work was supported by the FIS grant 94/0728 from the Spanish Government.

BIBLIOGRAPHY
1. Ashford, R.W., Desjeux, P., deRaadt, P. Estimation of population at risk of infection and number of cases of leishmaniasis. Parasitol Today 1992; 8: 104-105.
2. Ouellette, M., Papadopoulou. Mechanisms of drug resistance in Leishmania. Parasitol Today 1993; 9: 150-153.
3. Segovia, M. Leishmania gene amplification: A mechanism of drug resistance. Ann Trop Med Parasitol 1994; 88: 123-130.
4. Beverley, S.M. Gene amplification in Leishmania. Ann Rev Microbiol 1991; 45: 417-444.
5. Navarro, M., Maingon, R., Hamers, R., Segovia, M. Dynamics and size polymorphisms of minichromosomes in Leishmania major LV-561 cloned lines. Mol Biochem Parasitol 1992; 55: 65-74.
6. Ortiz, G., Navarro, M., Segovia, M. Location in the source chromo some of the 180-kb minichromosome of Leishmania major and characterization of the novel junction. Mol Biochem Parasitol 1995; 71: 153-161.
7. Coderre, J.A., Beverley, S.M., Schimke, R.T., Santi, D.V. Overproduction of a bifunctional thymidilate synthetase-dihydrofolate reductase and DNA amplification in methotrexate-resistant Leishmania tropica. Proc Natl Acad Sci USA 1983; 80: 2132-2136.

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