Insect Biochem., Vol. 10, pp. 535 to 541. 0020-1700/'80/1001-0535 $02.00/0
© Pergamon Press Ltd. 1980. Printed in Great Britain.
ONTOGENY AND TISSUE DISTRIBUTION OF LEUCINE
AMINOPEPTIDASE IN DROSOPHILA MELANOGASTER*
VIRGINIA K. WALKER~ and JOHN H. WILLIAMSON
Department of Biology, University of Calgary, Calgary, Alberta, Canada T2N 1N4
(Received 26 November 1979)
Abstract--Drosophila melanogaster larvae contain two isozymes of leucine aminopeptidase that are
detectable on starch gels. The more anodally migrating isozyme, LAP A, is first detected 12 hr after
oviposition and is a soluble enzyme localized to the haemolymph. The specific activity of the other isozyme,
LAP D, is highest in young larvae and associated with cell membranes, especially from the midgut and
Malpighian tubules. It is probable that these two enzymes are both functionally and spatially unique in
larvae of D. melanogaster.
Key Word Index: Leucine aminopeptidase, Drosophila melanogaster, zygotic gene expression, cell
fractionation, tissue specificity of enzymes
INTRODUCTION
DESPITE the presumptive physiological importance of
peptidases their biochemical analysis in insects is
fragmentary (LAW et al., 1977). The first report of
peptidases in Drosophila melanogaster was that of
BECKMAN and JOHNSON (1964). The two leucine
aminopeptidase (LAP) isozymes observed in third-
instar larvae (SAKAI et al., 1969) were found to be
products of two genes Lap-A and Lap-D. These genes,
on chromosome 3-position 98.3, are so closely linked
that no recombination between them has been
observed (BECKMAN and JOHNSON, 1964; FALKE and
MACINTYRE, 1966; LINDSLEYand GRELL, 1968). It has
been suggested that these enzymes are 'pupal enzymes'
which are responsible for the histolysis of larval tissues
(MUHS, 1975). Thus the presence of leucine
aminopeptidases in larval stages of development
assumes major interest. It is possible that these
histolytic enzymes are synthesized early in
development and stored in an inactive form prior to
utilization. Therefore, the subcellular localization and
tissue distribution of the LAP A and LAP D isozymes
is also of interest. In this paper a biochemical assay is
described for Drosophila LAP which was used to study
the developmental expression and the cellular
localization of this enzyme in Drosophila
melanogaster.
MATERIALS AND METHODS
Drosophila stocks
Wild type (Oregon-R) Drosophila melanogaster cultures
were reared at 19-21°C on standard cornmeal medium
(LEw~s, 1960). Eggs and embryos were collected on 3~ agar
*Supported by the National Sciences and Engineering
Research Council of Canada.
~'Present address: Department of Genetics, University of
Cambridge, Cambridge, CB2 3EH, U.K.
which was smeared with a yeast and vinegar paste containing
a few drops of alcohol to stimulate oviposition (MITchELL
and MXTCnELL, 1964). Unless specified, Oregon-R third-
instar larvae were used for all experiments.
Stocks homozygous for particular LAP electrophoretic
variants were produced by homozygosis of third
chromosomes from wild type stocks. Several lines from each
stock were subjected to electrophoretic analysis for
identification. A line homozygous for the fast variant of
Lap-D (Lap-DF) and the fast variant ofLap-A (Lap-AF) was
isolated from K~is 60. A line carrying the null allele for Lap-A
(Lap-A °) and Lap.DF was isolated from Kiis 51. To
determine the time of initial expression of the Lap-A gene,
Lap-A o Lap-DF virgin females were mated with Lap-AF
Lap-DF males, and progeny embryos were analyzed. To
facilitate the collection of large numbers of parents for
sp~ific crosses, stocks carrying C(1)RM, ts l(1)E-90 or
XyLys, ts 1(1)E-90 were constructed. This temperature
sensitive allele (kindly supplied by Dr. D. SuzuKI,
Vancouver, Canada) with appropriately timed heat shocks
(29°C) results in cultures of only one sex. Unfertilized eggs
from Lap-A ° Lap-DF females were obtained by crossing
virgins of this genotype with sterile X/O males.
Enzyme assays
Leucine aminopeptidase activity was measured by
modification of the method for the determination of rat brain
amidase activity (MARKS et aL, 1968). Samples were obtained
by homogenizing whole animals in distilled water and centrifug-
ing at 8000 g (4°C) for 3 rain. The assay solution contained 0.7
ml of 0.1 M Tris-maleate buffer, pH 6.7, and 0.3 ml of 1.8 mM
L-leucyl-fl-naphthlamide--HCl (Sigma Chemical Co. St. Louis,
MO, U.S.A.) in distilled water. The reaction was initiated by
addition of the supernatant and incubated at 30°C for 20 min.
Incubations were terminated by adding 0.5 ml of a solution
containing 5% Tween-20 and 0.1% Fast Garnet GBC salt in
1.0 M acetate buffer, pH 4.2. Colour was allowed to develop
for 10 rain at 22°C after which the samples were kept on ice
until their absorption was measured at 520 nm. Reference
blanks were prepared by the addition of the enzyme after
inclusion of the coupling dye. Standard curves were prepared
with fl-naphthylamine (flNA) (Sigma Chomcial Co.)
dissolved in 40% ethanol.
The sensitivity of the enzyme to the inhibitors EDTA and
1, 10-phenanthroline was tested by preincubating the enzyme
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