Cell Motility and the Cytoskeleton 37:300–307 (1997)

Patterns of Microtubule Assembly in

Taxol-Treated Early Drosophila Embryo

Giuliano Callaini and Maria Giovanna Riparbelli

Department of Evolutionary Biology, University of Siena, Siena, Italy

Incubation of early Drosophila embryos with low concentrations of taxol (2.5 µM)
revealed a pattern of microtubule assembly that was cell-cycle dependent.
Microtubule bundling was observed during the pronuclear stage after resumption
of meiosis, whereas at the onset of the first mitosis the microtubules organized in
astral arrays. Taxol treatment showed differential microtubule assembly properties
of the egg cytoplasm. The preferential assembly site for taxol-induced asters was
the ventral cortex; in the dorsal cortex only microtubule bundling occurred. This
dorsal-ventral heterogeneity of the egg cortex persisted until the third or fourth
nuclear cycle. Microtubules did not organize in astral arrays in the inner cytoplasm,
but only in mitotic spindles. CP190 and g-tubulin, usually found in the centrosome
of the early Drosophila embryo, were absent in taxol-induced asters. These
observations suggest that the mechanism driving the assembly of taxol-induced
asters is not centrosome dependent in the early Drosophila embryo. Cell Motil.
Cytoskeleton 37:300–307, 1997. r 1997 Wiley-Liss, Inc.

Key words: cytoplasmic heterogeneity; taxol-induced asters; mitosis

INTRODUCTION ences in the organization of the microtubule cytoskeleton

The early Drosophila embryo is a large cell in
which 13 mitoses occur without the formation of distinct
membranes between neighbouring nuclei. The dividing
nuclei remain in the interior of the embryo until the tenth
mitosis when they migrate to the cortical region and
divide three times more before cell membranes are
formed [Foe et al., 1993]. Several studies suggest that the
cytoskeleton plays an important role in early develop-
ment [Warn, 1986; Schejter and Wieschaus, 1993; Foe et
al., 1993; Miller, 1995]. Microtubules seem to form a
filamentous network just beneath the plasma membrane
of the preblastoderm embryo [Karr and Alberts, 1986].
This cortical network is not, however, visible in embryos
fixed without taxol [Warn and Warn, 1986; Kellogg et al.,
1988; Baker et al., 1993]. This finding suggests that
peripheral microtubules could be an artifact of taxol
treatment. However, short randomly oriented microtu-
bules occur throughout the oocyte before ovulation
[Theurkauf et al., 1992].
Although the egg has established its principal axes,
dorsal-ventral and anterior-posterior, by the end of oogen-
esis [Chasan and Anderson, 1993], no regional differ-
are apparent in the early embryo. The dorsal-ventral
polarity of the early embryo can only be predicted from
the shape of the egg that is curved on the ventral side and
flattened on the dorsal one. Dorsal-ventral asymmetry
first becomes apparent at the end of cellularization, the
ventral cells being completed before the dorsal cells
[Sonnenblick, 1950]. To test for cytoplasmic heterogene-
ity in microtubule assembly properties we treated early
Drosophila embryos with low concentration of taxols.
Taxol is a plant-derived drug that has been shown to
inhibit cell division [Wani et al., 1971] and to promote
microtubule assembly by lowering the critical concentra-
tion required for tubulin polymerization [Schiff et al.,
1979] in a cell-cycle dependent manner [De Brabander et
al., 1986]. By using this drug we obtained evidence of
Contract grant sponsor: MURST.
*Correspondence to Dr. Giuliano Callaini, University of Siena, Depart-
ment of Evolutionary Biology, via Mattioli, 4, 53100 Siena, Italy.
E-mail: callaini@unisi.it
Received 23 August 1996; accepted 27 March 1997.

r 1997 Wiley-Liss, Inc.


cytoplasmic domains with different microtubule assem-
bly properties in the preblastoderm embryo. This informa-
tion provided insight into the cytoskeletal organization
and dynamics of early Drosophila embryogenesis.
MATERIALS AND METHODS
Reagents
Microtubules were detected with a monoclonal
antibody against b-tubulin (Boehringer Mannheim, India-
napolis, IN) and a monoclonal antibody against acety-
lated a-tubulin (Sigma, St. Louis, MO) both used at
1:200-fold dilution. The centrosome-associated protein
CP190 was detected with the rabbit polyclonal antiserum
Rb188 [Frasch et al., 1986; Oegema et al., 1995; Whit-
field et al., 1988, 1995] used at 1:400 dilution; g-tubulin
was detected using Rbcsl, a polyclonal antiserum raised
against recombinant Drosophila g-tubulin (dilution,
1:100). The antibodies against g-tubulin and centrosomal
antigens of Drosophila were kindly provided by W.G.F.
Whitfield, University of Dundee, United Kingdom. Sec-
ondary antibodies were either goat anti-mouse or goat
anti-rabbit conjugated IgG (Cappel, West Chester, PA;
dilution, 1:600) coupled with fluorescein or rhodamine.
Nuclei were visualized with 1 µg/ml of the specific dye
Hoechst 33258 (Sigma). Taxol (Molecular Probes, Eu-
gene, OR) was dissolved in dimethylsulfoxide (DMSO)
as stock solution and diluted with D20 medium [Echalier
and Ohanessian, 1970] to the final concentration desired
for incubation. Bovine serum albumin (BSA) was ob-
tained from Sigma.
Egg Collection
Drosophila melanogaster (Oregon-R) flies were
raised in groups of 50 males and 25 females on standard
medium in 200-ml plastic containers. Eggs from 7- to
8-day-old flies were collected twice for 20 min on small
agar plates at 24°C. After discarding these eggs, fertilized
eggs were collected again at 5-min intervals.
Drug Treatment
For taxol incubation, the eggs were dechorionated
in a 50% bleach solution, washed in distilled water, dried
on filter paper, and permeabilized with heptane on small
Petri dishes for 2 min as described by Limbourg and
Zalokar [1973]. The eggs were then quickly dried,
covered with a small square of tissue paper (Kleenex),
and incubated with the drug for 15 min at 24°C. The
tissue paper prevented the eggs from floating on the
surface of the solution and ensured that the whole embryo
was in contact with the drug.
Controls were performed by incubating the eggs
without taxol in D20 medium containing 0.5% DMSO.
Microtubules in Taxol-Treated Drosophila Embryo 301
DMSO at a concentration of less than 1% does not
interfere with embryo development [Zalokar and Erk,
1976].
Fluorescence Microscopy
To remove the vitelline envelope, the eggs were put
in a solution of 5 ml 90% cold methanol in H2O plus 5 ml
n-heptane and shaken for 4–5 min (Warn and Warn,
1986). Eggs that lost their vitelline envelope sink down.
These eggs were fixed for 10 min in methanol and for 5
min in acetone, both at 220°C. After fixation, the eggs
were washed in phosphate buffered saline (PBS) and
incubated for 30 min in PBS containing 0.1% BSA. To
visualize CP190 antigen or g-tubulin, the embryos were
incubated with Rb188 or Rbcs1 antibodies at room
temperature. After 4–5 h, an antibody against b-tubulin
was added for 2 h. The embryos were then rinsed in
PBS/BSA and incubated for 1 h in a mixture of secondary
antibodies. To exactly determine the stage of the em-
bryos, the nuclei were stained with Hoechst 33258 for
3–4 min. The samples were rinsed again in PBS and
mounted on glass microscope slides in 90% glycerol
containing 2.5% n-propyl gallate [Giloh and Sedat,
1982]. Fluorescence observations were carried out with a
Leitz Aristoplan microscope equipped with fluorescein,
rhodamine, and UV filters. Photomicrographs were taken
with Kodak Tri-X 400 pro and developed in Kodak
HC110 developer for 7 min at 20°C.
RESULTS
Taxol Induced Cell-Cycle Dependent Patterns
of Microtubule Assembly in the Early
Drosophila Embryo
The early Drosophila embryo has a thin peripheral
layer that was faintly stained by an antibody against
b-tubulin without revealing distinct microtubule bundles
(not shown). Only microtubules associated with the
meiotic apparatus or with polar bodies were recognized.
The inner cytoplasm was dark and only the spindle
microtubules were labelled.
Incubation with taxol (2.5 µM for 15 min), an agent
known to promote microtubule assembly by lowering the
critical concentration required for tubulin polymerization,
led to a dramatic reorganization of the peripheral cytoplas-
mic domain and induced the formation of distinct micro-
tubule bundles. By immunofluorescence with an antibody
against b-tubulin, we observed a complex pattern of
microtubule organization that was correlated with the
behavior of the parental complements. When embryos
collected from rapidly laying females were incubated in
taxol within 5 min of egg deposition and scored for DNA,
we found three main nuclear configuration types: pro-
302 Callaini and Riparbelli
nuclear migration stages (19%, n 5 196), first mitotic
figures (70%, n 5 682), and late nuclear divisions (11%,
n 5 107). This variety may be due to the fact that it is
difficult to obtain synchronized batches of eggs from
Drosophila females. Our attempts to obtain the various
meiotic stages were unsuccessful. Because the time
required to complete meiosis in Drosophila was shorter
than the time of taxol treatment, the lack of meiotic
figures suggests that taxol, at a concentration of 2.5 µM,
did not significantly affect meiotic progression in Dro-
sophila. One hundred sixty-nine eggs of the first class
showed four peripheral nuclei, sometimes arranged in
tandem, and a single nucleus in the inner cytoplasm. This
finding indicated that meiosis had been completed but
that pronuclear migration had not occurred in these
embryos. Heterogeneous bundling of microtubules, rang-
ing from parallel bundles to partial clustering, was
induced by taxol under these conditions (Fig. 1A and B).
In 27 eggs of the first class, the male and female pronuclei
were closely apposed in prophase-like configuration.
Cortical bundles of microtubules, mostly parallel to the
surface, were seen to converge toward bright foci in these
embryos (Fig. 1C and D). Microtubule bundles persisted
during prometaphase, but decreased in number. From
metaphase of the first mitosis, the microtubule bundles
were no longer visible, or only in the dorsal region of the
embryo, and loosely assembled asters appeared in the
ventral cortex (Fig. 1E and F).
When the embryos were incubated with D20 me-
dium without taxol and containing 0.5% DMSO, we
never observed astral arrays or microtubule bundles at the
egg surface. Only microtubules associated with polar
bodies or mitotic spindles were visible (Fig. 2).
As nuclear division progressed, the surface of the
embryos treated with taxol was filled with distinct
aster-like configurations connected by a network of short
microtubules (Fig. 3A and B). With a higher concentra-
tion of taxol (10 µm), the microtubule asters increased in
size and compactness and took the shape of hollow
spheres with walls formed by short hook-like radially
orientated microtubule bundles (Fig. 3C and D). With
longer incubation times the asters eventually merged.
This finding was suspected because paired asters and
large masses of radially oriented microtubules were
found (not shown).
Taxol Incubation Revealed Regional Differences in
the Early Drosophila Embryo
The pattern of taxol-induced microtubule polymer-
ization revealed a dorsal-ventral cytoplasmic heterogene-
ity of the egg cortex. The organization of the microtu-
bules in astral arrays appeared to be an intrinsic property
of the ventral region of the embryo, whereas the dorsal
one showed only randomly arranged microtubules (Fig.
1E and F). The asters decreased in size and number along
the transition zone: the dorsolateral area of the egg, where
short microtubules were also visible. The dorsal-ventral
cytoplasmic heterogeneity was no longer visible from the
third or fourth nuclear cycle.
Taxol-induced microtubule bundling was appar-
ently a specific property of the peripheral cytoplasm,
because we never observed asters or microtubule bundles
in the embryo in addition to the spindle microtubules
(Fig. 3E). Taxol at a concentration of 2.5 µM, induced
dramatic reorganization of the cortical microtubules,
without noticeably affecting the spatial organization of
the spindle microtubules (Fig. 3E). This failure to affect
microtubule organization in the embryo was not due to
lack of taxol penetration, because spindle organization
was unaltered in eggs cut longitudinally and in syncytial
blastoderms, in which the nuclei had migrated to the
embryo surface. Abnormally large asters were induced at
the spindle poles at a higher concentration of taxol (10
µM). The asters produced by adding taxol were always
visible at the embryo surface during mitoses 1–9, but
were no longer observed as soon as the nuclei reached
the periphery of the embryo during the tenth mitosis
(Fig. 3F).
Asters Induced by Taxol Differed From the Asters
at the Spindle Poles
Cortical asters did not seem to contain acetylated
a-tubulin, because we were unable to obtain surface
labeling in embryos treated with 2.5 µM taxol (Fig. 4A
and B). This finding was not due to lack of antibody
penetration because, as a rule, acetylated tubulin was
found in polar bodies (Fig. 4A) and throughout the
mitotic spindles (Fig. 4B) in the interior of the embryo.
Acetylated microtubules seemed to be associated with a
thin area through the longitudinal axis of the mitotic
spindle, whereas acetylated a-tubulin was found in asters
assembled at higher concentration of taxol (10–20 µM)
(not shown).
Rb188 antibody that recognized a protein of 190
kDa associated with the centrosome of the early Dro-
sophila embryo did not stain the central area of taxol-
induced asters, but recognized the CP190 antigen at the
poles of the mitotic spindles in the inner cytoplasm.
Figure 4C and D shows an example of centrosome
staining of the spindle poles obtained with the Rb188
antibody. Although g-tubulin was found at the poles of
the spindles in the inner cytoplasm, it was not found in the
cortical asters (not shown).

Fig. 1. Drosophila embryos treated with 2.5 µM taxol for 15 min and
stained with an antibody against b-tubulin. A,B: At the end of meiosis
the embryo surface is filled with short randomly orientated microtu-
bules. C,D: When parental nuclei are in close apposition, the periphery
of the embryo shows bundles of microtubules mostly converging to
common foci (arrowheads). E,F: During metaphase of the first mitosis,
Microtubules in Taxol-Treated Drosophila Embryo 303
small astral arrays of microtubules are evident in the ventral and lateral
regions of the egg cortex; the dorsal region shows short randomly
orientated microtubules. Arrows indicate polar bodies. Embryo in F has
a right-angle clockwise orientation with respect to that shown in E.
Bars 5 100 µm in A,C,E, and 10 µm in B,D,F.
304 Callaini and Riparbelli
Fig. 2. Microtubule distribution in Drosophila embryos incubated for
15 min in culture medium containing 0.5% DMSO without taxol. Note
that only microtubules of the polar bodies are visible (arrows). Astral
arrays or microtubule bundles are absent. The embryo in the middle is
at the same stage as that in Figure 1C; the embryo on the left is at the
second nuclear cycle; the embryo on the right corner is at the stage of
pronuclear fusion. Bar 5 100 µm.
DISCUSSION
In line with earlier observations in PtK2 cells
[De Brabander et al., 1986] and Xenopus egg cell-free
extracts [Verde et al., 1991], taxol treatment induced
microtubule assembly in a cell-cycle-dependent manner
in the early Drosophila embryo. We observed cortical
microtubule bundles during pronuclear stages, and dis-
crete astral arrays of microtubules during the first mitosis.
In asters induced by taxol treatment, the lack of
centrosomal components such as the CP190 antigen
[Whitfield et al., 1988, 1995] and g-tubulin [Sunkel et al.,
1995], typically found at the spindle poles of the mitotic
apparatus of the early Drosophila embryo, suggests that
the mechanism of assembly of the asters differs from that
required for spindle pole organization. This theory is
consistent with the hypothesis of centrosome-indepen-
dent assembly of microtubules in radial structures in the
presence of taxol observed in vivo and in vitro [Schatten
et al., 1982; Maekawa and Kuriyama, 1991; Verde et al.,
1991; Kallajoki et al., 1992; Gaglio et al., 1995]. The
morphology of taxol-induced asters is also indicative of
an unusual assembly mechanism. These asters are hollow
spheres of different sizes without a central core or distinct
focus for radial microtubules. The latter were closely
packed to form the wall of these structures. Only microtu-
bules of the asters observed during the first mitosis
seemed to converge to a central core. This finding,
however, may be due to the fact that these asters are so
small that their structure cannot be resolved. Verde et al.
[1991] demonstrated that cytoplasmic dynein was re-
quired for aster assembly in taxol-treated extracts of
Xenopus eggs, because the mitotic asters failed to orga-
nize correctly in the presence of 10 µM of vanadate.
During the early preblastoderm stages, we found
that microtubule assembly did not occur in a homoge-
neous manner through the cytoplasm. We observed that
many astral arrays of microtubules organized in the
ventral periplasm in the presence of taxol (2.5 µM),
whereas microtubule bundling was irregular in the dorsal
region. This cytoplasmic asymmetry in microtubule dy-
namics is intriguing because it predicts dorsal-ventral
polarity, before organizational differences with respect to
this axis were apparent in the embryo. Asymmetry along
the dorsal-ventral axis of the Drosophila embryo first
becomes evident at the end of cellularization when
ventral blastoderm cells are completed before the dorsal
cells [Sonnenblick, 1950]. Regional differences in aster
microtubule organization have also been described during
early Beroe development when taxol induced the assem-
bly of small asters around the mitotic spindle and
bundling of microtubules in the opposite egg cytoplasm
[Houliston et al., 1993]. In Drosophila, the astral arrays
of microtubules gradually filled the embryo surface, and
regional heterogeneity disappeared when more nuclei
populated the interior of the embryo during the fourth
mitosis. Interestingly, taxol did not induce microtubule
polymerization in the interior of the embryo, where
spindle microtubules assembled. This finding suggests
that tubulin concentrations in the inner cytoplasmic
domain do not warrant taxol-dependent microtubule
assembly, but only allow spindle microtubule nucleation
from the centrosomes.
Taxol at a low concentration induced microtubule
organization in cortical asters, but not accumulation of
detectable levels of acetylated a-tubulin in the preblasto-
derm embryo. The lack of acetylated tubulin in taxol
induced asters is apparently in contrast with previous
findings showing bright staining of cortical microtubule
clusters in taxol-treated embryos [Wolf et al., 1988]. This
discrepancy may be due to the lower taxol concentration
(2.5 µM) and to the shorter incubation times (15 min) we
used in our experiments. We observe faint labeling of the
mitotic spindles. This finding suggests that the accumula-
tion of acetylated tubulin required different taxol concen-
trations and/or different exposure times between the inner
cytoplasm and the cortex of the preblastoderm Dro-
sophila embryo. At the concentration used (2.5 µM), taxol
did not significantly affect astral microtubule organiza-
tion at the spindle poles, whereas higher concentrations

Fig. 3. Microtubule staining of Drosophila embryos treated for 15 min
with 2.5 µM (A,B,E,F) or 10 µM (C,D) taxol. A,B: Anaphase of the
fourth nuclear cycle; the egg cortex is filled with astral structures
connected by short microtubules. Note that many of the astral arrays
have an empty core (arrows). Anaphase spindles appear in a lower
focal plane. C,D: Embryos during metaphase of the third nuclear
division cycle treated with a higher taxol concentration. Spherical
aggregates of short microtubules covered the entire surface of the
Microtubules in Taxol-Treated Drosophila Embryo 305
embryo. Cross-view of the microtubule aggregates show an empty
core. E: Cross-view of an embryo during anaphase of the eighth
mitosis; taxol-induced asters are visible in the periplasm (arrows), not
in the inner cytoplasm. Note the normal-shaped mitotic spindles. F:
Surface view of an embryo during prophase of the tenth mitosis; note
the lack of taxol-induced asters. Bars 5 100 µm in A,C, 10 µm in B,D,
and 40 µm in E,F.
306 Callaini and Riparbelli

Fig. 4. Drosophila embryos treated with 2.5 µM taxol for 15 min. (A) Surface and (B) cross-views of an
embryo stained with an antibody against acetylated a-tubulin during prophase of the third mitosis. Only a
few microtubules associated with polar bodies (arrowhead) and with the mitotic spindles (arrows) are
acetylated. (C) Tubulin and (D) Rb188 labeling in an embryo during prophase of the eighth mitosis;
centrosomal material (arrowheads) was found near the spindle poles (arrows), not close to taxol-induced
asters (empty arrows). Bars 5 40 µm in A,B,C, and D.

disrupted spindle architecture and produced a large
amount of astral microtubules [Wolf et al., 1988].
ACKNOWLEDGMENTS
We thank W.G.F. Whitfield for the generous gift of
Rbcs1 and Rb188 antibodies. This work was supported
by MURST (40 and 60% grants).
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