Contributions
Zoology,
to
SPB Academic
Chromosomal
for
Barranco J. Cabrero
P.
de
Departamento
E-04I20 La
bilateral
a
Escuela Politécnica
Biología Aplicada,
2
Cañada, Almería, Spain;
Universidad de
Keywords:
3
Departamento
Departamento
E-18071
Granada,
de
de
Granada,
Abstract
left side
in
was
bilateral
and
the
than
and the
as
Sierra
side
of the
had
and
a
spontaneous
in
shown
with
natural
a
that the
almost
ovipositor
dorsal valves
specimen
The
have
female,
valves
a
found
Nevada,
side
right
two spurs;
normal.
the left
on
of
analyses
Pycnogaster inermis,
Spanish
male
short, appearing
testis
cytogenetic
of
symmetry. Ventral
shorter
Animal
Biología
were
well-developed
very
longer but
apparently
an
perfect
were
ovary
y
en
Tettigoniidae,
los individuos
do
en
El
origen
la formación
de
espermatozoides,
explicación
un
más
óvulo binucleado
cigoto hembra (XX)
en
la
chromosomes in
the testis
the presence of 2n
and 2n
28
=
=
+ XX in the
28
as
standard
cies.
This
very
early
have
originated
males
that
suggests
and
the
developmental
from
un
X
cromosoma
división de la
la
en
segmentación.
ovary, the
gynandromorphism
one
This
stage.
formation
the
male-
of this
females, respectively,
second-polar-body reactivation
two spermatozoa,
is
and
binucleate
a
subsequent
and the
occurred
other
at
a
could
gynandromorph
of
however,
one
first
the most
egg
by
fertilization
by
nomenon
among
reported
in
Chopard
(1979)
is the
have
in a female-determined
at the
zygote (XX)
most
the
cases,
external
Resumen
ción
natural de Sierra
macho y el lado
perfecta.
que las
muy
de
Las
una
cortas, casi
El
análisis
cromosomas
en
como
el
normal
lado
en el
del
que el
con una
lado
eran
cortas
las ventrales
izquierdo
y un
un
eran
testícu-
ovario
reveló
y
la
presencia
=
28
+
re-
de 2n
XX
en
Orthoptera.
=
el
limited
was
(1979)
in
reported
which
he
had
noted,
received
a
analysis
of male and
a
In
study
reviewed
been
few
100
over
found
a
of
sometimes
Orthoptera,
only
full examination
to
and
genitalia,
Hewitt
dromorphs
in
about
Tettihave
cases
chromosome
including
female tissues.
show bilateral symmetry,
male and the other female,
terns
Most gynan-
other
have
also
been
28
one
although
showing predominance
Differing patterns
de 2n
in
of
described
one
halfbeing
several
sex
over
(Hewitt,
patthe
1979).
bien
derecho.
el testículo
and
(1966),
comprehensive
izquierdo
más
Internamente había
lado
the
including
pobla-
en una
simetría bilateral
oviscapto
mientras que
espolones.
citogenético
en
hembra,
valvas dorsales
espécimen ginan-
encontrado
Nevada mostraron
derecho
hembra normal
aparentemente
desarrollado
inermis
un
of
As
gonioidea.
Pycnogaster
published
report
organs.
quarter
one
de
and has been
1965)
Uvarov
(1938),
morphology,
gynandromorphs
dromórfico
phe-
common
elimination
division.
morfológicos y citogenéticos de
(Crew,
gynandromorphism
on
internal
Los análisis
relatively
different groups
many
Orthoptera.
Hewitt
insects
female-determining;
parsimonious explanation
X chromosome
cleavage
a
spe-
views
lo
femenino; pero
y otro
de
del
éste por
X0
+
Gynandromorphism
same
casi
primera
estar basa-
por reactivación
subsiguiente de
la eliminación
sugiere
Introduction
revealed
Cytogenetic analysis
era
fertilización
masculino
uno
simple es
Esto
especie.
estadío muy temprano
right
side.
of
de la
un
ginandromorfopudo
de este
un
segundo corpúsculo polar y
dos
normales
ginandromorfismo se generó en
desarrollo.
del
Orthoptera
normal
the
on
still
Universidad de
Facultad de
Ecología,
Spain
mismo que
gynandromorph
Universidad de Almería,
Superior,
que el
population
(1995)
Amsterdam
Pycnogaster inermis
Genética, Facultad de Ciencias,
cytogenetics, Pycnogaster inermis,
Gynandromorphism,
Morphological
123-127
J.P.M. Camacho & F. Pascual
Granada, E- 18071 Granada, Spain;
Ciencias,
in
gynandromorph
(2)
1838) (Orthoptera, Tettigoniidae)
(Rambur,
1
basis
65
Publishing bv,
+
X0
ovario,
lo
of
gynandromorphism
sible in insects because the
cell
individually,
so
sex
that the
are
fea-
is determined in each
genetic
constitution of
P. Barranco
124
et
al.
for
Chromosomal basis
a
female tissues in the adult
on
the
moment
lost
wards,
a
clear
if
Thus,
the second
during
gynandromorph depends
which the X chromosome is lost
at
1992).
(Lawrence,
gynandromorph
bilateral
X
one
cleavage
chromosome is
division,
female
predominance of
tissues will follow. On the other hand, the
of male
nance
(see
Hewitt,
tissues,
could easily
1979),
non-disjunction
of the
male zygote,
a
in
in
reported
or
predomiacridoids
some
explained
be
from the second
division onwards, giving rise
male
by
X chromosome of
single
stage
after-
over
to a
a
cleavage
proportion of
XX
cells lower than 50%.
In
the
Orthoptera,
netic marker
evidence
provided
nism
in
only reported
(which produced
the
for
a
grasshopper
a
Valanga
1838).
and
Morphological
fect bilateral
of
nance
the
over
1-3. Posterior part of the
Figs.
gynandromorph, (2)
Scale
bar
=
0.5
a
normal
abdomen
male
(dorsal view)
and
(3)
a
of
normal
XX.
were
of this
specimen
predomi-
no
and that
XO,
were
Hypotheses
analyses
almost per-
an
with
other,
spermatocytes in the testis
gin
was
but
primary
ovariole-
explain
to
of the
(Rambur,
cytogenetic
specimen
gynandromorph,
one sex
wall cells
irregularis
inermis
Pycnogaster
have shown that this
ge-
The present paper
spontaneous gynandromorph
tettigonid species
a
mosaicism),
the binucleate egg mecha-
(Walker, 1870) (cf. White, 1968).
deals with
with
case
colour
the ori-
discussed.
are
(1) the
female.
cm.
Materials and methods
each cell determines its
for
The
sex.
own
system (White,
Of the several
Orthoptera
is
sex-deterOne
the X0 cr/
1973).
cytogenetic
bilateral
explain
expression
predominant chromosomal
mination mechanism in
XX 9
phenotypic
to
theories advanced
gynandromorphism,
Orthoptera: (1)
a
one
to
apply
two
dinates:
the
one
of the nuclei is fertilized
sperm and the other
by
the X chromosomes in
at
the first
cleavage
morphology,
between
0 sperm; and
an
an
by
(2)
an
one
X
of
XX zygote is eliminated
division.
these
To
distinguish
ac-
female
tion.
two
hypotheses,
are
essential.
Such
markers
that the second mechanism is the
in
Drosophila,
and that the
more
proportion
have
fixed for
and the
with 0.05%
at
a
of
The
del Chullo
2280
detailed
were
internal
colchicine
gonads
acid and stored in
anaesthesized
anatomy,
orcein. To
then immersed
refrigerator at
perform
the
into
4°C. For
by squashing
the
and
For this
in
dissected
which
purpose,
the
the
the
abdomen,
prior to
3:1
coor-
of external
after
in insect saline solution 6 h
made
UTM
examination
cytological analysis.
were
a
(Sierra Nevada,
altitude;
m
gynandromorph were injected,
preparations were
dissec-
ethanol:acetic
cytological analysis,
different materials
in
C-banding technique, squashing
genetic
was
markers
gynandromorph of Pycnogaster
Morrón
the individuals
thorough study
a
acetic
curately
at
binucleate egg is produced
failure of the second meiotic division and,
subsequently,
and one
30SWG0206. After
gonads were
by
female,
collected
were
Huéneja, Granada, Spain)
for
best
male,
inermis
shown
frequent
one
of male and
done
in 45% acetic acid and the
immersing
was
al.
the
preparation
performed following
(1984).
in
the
coverslip
was
liquid nitrogen.
removed
The
technique described
after
C-banding
in Camacho et
65
Zoology,
to
Contributions
(2)
125
1995
-
Results
Morphological analysis
The
general
the
gynandromorph
normal
specimens
(Figs.
1-6).
structures
ones
The
were
of
the left tegmen
showed
on
was
sinuous
a
body
but the
1 -2). The soft
to
more
venation
was
structures
hiding
and
irregular
(Fig. 4).
male features
were
wider than normal
the titillator
rounded apex. The
cer-
(Figs.
of the male genitalia
titillator.
apThe
gynandromorph.
single
a
that of males. Ab-
as
were
Although
small, flat
was
subgenital plate
deformed, reduced and slightly
the
was
situated normal male
was
sclerotized
completely
a
properly
epiproct
conspicuous,
and with
External
-
a
ventral
halves.
sex
size similar
superficial
the left side of the
had
specimen
cus,
a
Tegmina
suture closed in the middle
parent
but the
in both
females, but the internal edge of
dominalsternites showed
Male features.
of
female fea-
nor
formed,
females.
that in
to
of
dif-
clearly
were
normal male, but the pronotum
characteristic
similar
that
The dorsal terminal abdomiwell
gynandromorph
a
shape)
to
but the external
species,
equally amorphous
were
that of
similar
Neither male
predominated.
tures
nal
of this
and
size,
rather
was
and abdominal sternites
genitalia
ferent
appearance (colour,
was
outside
displaced
Figs.
body (Fig. 4).
There
was
apparently
an
ternal left side.
seminal vesicles
the accessory
Regarding
were
well
normal testis in
developed
glands,
situated
ejaculatorius
on
was
the
right body
not
the
the
(a
4-6.
=
sclerites
half. The
which
=
e
0.5
had
on
its
right
=
right
smaller than
dorsal
a
cercus
was
normal and
Figs.
were
quite
and
1
3).
a
reduced
small
the
to a
highly
female
valves
were
=
left dorsal
right
valve).
ventral
Scale
bar
one
and
half the
showing
rounded
a
twisted
being slightly
length
of the
dorsal
right
right body
ovary full of eggs
half there
at
was
a
well-
differentmaturation
however,
subgenital plate
callus situated
(Fig. 4).
valves, extremely short,
f
side. The fe-
The
were
longer,
reduced
the
close
ovipositor
four deformed valves of differentsizes.
Dorsal
=
stages. The oviduct and the spermatheca
were
ap-
normal.
sclerotized lateral lobe and
rough amorphous
left ventral valve
(4)
sclerite of the
=
d
of
normal male
(Figs. 4—6).
parently
was
b
valve,
a
normally positioned
Ventral structures,
modified. The
(6)
developed
developed
(cf.
valve,
normal
On the internal
male
subgenital plate,
left ventral
and the left dorsal valve
measuring
valve
the external female features
=
(ventral view)
and
cm.
apex,
specimen
c
abdomen
normal female
ductus
interconnected with the ovi-
The
a
of the female
subgenital plate,
valve,
and
-
of the
part
and located on
duct.
Female features.
Posterior
gynandromorph, (5)
male
the left side and extended under the ovary,
was
the in-
Cytological analysis
The ventral
to two
right
to
had
one
spurs.
being
Conventional
that
staining
ovariole-wall cells
with
acetic
analyzed
orcein
showed
from the ovary,
P. Barranco
126
et
al.
-
Chromosomal basis for
The
of
application
chromatin
the
mining
provided
C-banding
of this
origin
for any hetero-
to test
that could
polymorphism,
exact
gynandromorph
gynandromorphism,
additional information,
no
in deter-
help
no
variations
C-banding pattern being observed, neither
in the
gynandromorph
the
bilateral
a
in
nor
the
in
normal indi-
two
viduals.
Discussion
The
the
fact that
specimen analyzed
was
that
the
early
in ontogeny. The
by
the
in
plain
of
the
cells
division. The
bilateral
with
one
and
without,
loss of
from
mechanism
gynandromorphy
that
as
ferti-
X chromo-
an
(2)
resulting
first
resulting
was
female zygote,
a
very
likely explanations
binucleate egg
a
spermatozoa,
one
cleavage
occurred
anomaly
X chromosomes in
two
ring
two
and the other
some
perfect indicates
polar body activation, which
from second
lized
almost
two most
derivation from
(1)
are:
developmental
of
gynandromorphism
the bilateral
one
of
occur-
the
first
could
ex-
reported by
White (1968).
the
Although
not
be ruled
XX zygote lost
binucleate egg
ties would be
Figs.
Mitotic
mosomes in
the
morph (Fig. 7)
cated
on
the
metaphase cells
ovary
located
and 2n
=
left side
28
on
right
28 +
=
side of the
+ XO chromosomes in
of the
body (Fig. 8).
of two chromosomes.
overlapping
showing
the
2n
XX chro-
gynandro-
the testis lo-
Asterisks
cause,
part
in
resulting
from the first
bar
5
=
body,
ber,
on
on
the
2n
28
=
+
typical
XX
female chromosome
(Fig. 7),
the left half possessed 2n
somes
(Fig. 8),
as
in the
testes
28
of
+
a
This chromosome number coincides
ported
by
Sentís
et
al.
(1988).
differences showed by the
fined halves of
the
mosomal
the
basis,
num-
and the testis located
=
two
normal male.
with that
the
had
possessing
chromosomes and the male half showing
re-
sexual
longitudinally
gynandromorph
female half
XO chromo-
Thus,
be-
organisms
would show, the male
nation would be XO instead of XY.
an
X elimi-
in fact
as
most
In XOcr/XXÇ
orthopterans
are,
cyto-
µm.
right side of the gynandromorph's
showed the
di-
possibili-
indicate
logical
situated
XY a/XX 9
an
one
cleavage
two
gynandromorph produced by
a
organisms,
Scale
in
easy
cytological analysis
as
is that
of the X chromosomes in
vision. The distinction between these
7-8.
can-
gynandromorph
simplest explanation
one
cells
two
a
in the P. inermis
the
reported here,
of the
hypothesis of
out
a
de-
chro-
two
only
X
one.
does
study
binucleate egg
which
We
purpose
have
means
of
Despite
reported
tried
the
markers
genetic
obtain
but
C-banding,
in
large
two
Orthoptera,
has been
partial
Geer,
1773).
for
made
a
1979).
by
In these
In
were
cytologi-
tettigoniids,
(1929)
in
Amblycorypha
and A.
as
by
gynandromorphs
few
Pearson
cases,
markers
unsuccessful.
were
only
1862)
indispensable.
cytogenetic
gynandromorphs,
tundifolia (Scudder,
are
number of
cally analyzed (Hewitt,
analysis
the
X-elimination hypotheses,
and
to
between
distinguish
not
this
only
ro-
oblongifolia (De
in P.
inermis,
the
Contributions
ovarian tissue showed
testicular tissue had
It
has
single
embryo (Houillon,
such
random and the
divide,
nuclei
resulting
first
do
in
that
the
of
cleavage
mix
completely
not
and
1992).
could be the
tera
male
or
A
the
as
Camacho, J.P.M.,
morphs.
tendency
Orthop-
to
become
gynandroreviewed
(1979)
the left half
was
termedia
female.
(1986) reported
1839)
a case
in
Platycleis
in which the left half
In the present P.
dromorph,
inwas
inermis bilateral gynan-
the left half was
number of left- and
would be
expected
by
(Hewitt,
these
a
1979).
the
a
random first
1938.
side
is
groups
as
In
fe-
always
could be
1984. C-
of
an
seg-
euchromatic extra
seg-
167-175.
La
biologie
des
1-291
Orthoptères:
(Le
(Methuen
&
1-188
(4th éd., revised):
Co., London).
1979.
G.M.,
and
Grasshoppers
3: Insecta
1
Orthoptera:
crickets.
In:
Animal
Born-
(Gebriider
1-170
Berlin/Stuttgart).
traeger,
1972.
Houillon, Ch.,
Embriología:
(Ediciones Omega,
1-184
Barcelona).
1992. The
Lawrence, P.A.,
making
of
a
fly:
1-228
(Blackwell
Publications, Oxford).
due
cleis
Martín,
J. & M.V.
goniidae). Actas
(Universidad
1929. The structure
Pearson, N.E.,
de
Diego,
VIII Jornadas Asociación
335-341
mología:
Peinado
1986.
Un
Platy-
(Serv.) ginandromórfico (Orthoptera,
intermedia
de
Española
Tetti-
de Ento-
Sevilla, Sevilla).
and chromosomes
J.
gynandromorphic katydids (Amblycorypha).
of three
Morph.,
47:
to
531-553.
period encompassed
embryo (Hewitt,
detection
1965. Sex-determination
Crew, F.A.E.,
Mateos
cases,
cleavage.
by
divisions of the egg and first cleavage
of the
Cabrero,
balanced
female
The different patterns shown
orthopteran
two
differences in
otic
male, giving
right-side
however, the right
Acridoidea,
male
from
J. Navas & J.
Chevalier, Paris).
Scientific
(3:3)
53:
Heredity,
cytogenetics
(Serville,
Viseras,
male
and, in the other two, female. Furthermore, Mateos
& Peinado
E.
grasshoppers:
Chopard, L.,
Hewitt,
of which
two
de
gynandromorphs
Hewitt
tettigoniids,
In
Andaluz
content of supernumerary chromosome
well-separated
female for each half of bilateral
four cases, in
the Plan
4068).
References
heterochromatin
of this feature in
test
analysis of
and
is
ment.
(Lawrence,
3122
of
ments of
male and female territories in the
and
(PB93-1108)
no.
first
organisms,
some
plane
large
D.G.I.C.Y.T.
Spanish
Investigación (Grupos
In
1972).
the
Drosophila,
as
they
127
X.
established
clearly
1995
-
X chromosomes and the
two
a
(2)
division determines bilateral symmetry
cleavage
the
been
65
Zoology,
to
the mei-
division
Sentís, C.,
and
genesis
1979).
Santos
J.
secondary
in
the
Uvarov, B.P.,
&
J.
nucleolar
genus
1966.
Fernández-Piqueras,
1988.
organiser regions during
Pycnogaster. Heredity,
Grasshoppers
and
locusts
60:
I:
Primary
spermato-
197-204.
1—481
(Cam-
bridge University Press, Cambridge).
White,
Acknowledgements
M.J.D.,
duced
by
White, M.J.D.,
We would
like
to thank
during the collection
morph was found,
tive
criticism.
Lola
of the
Campos-Carmona
samples
for her
among which this
help
1-961
1968.
double
study
was
supported by
grants
1973. Animal
gynandro-
from
the
Received:
gynandromorphic grasshopper
cytology
J.
Zool.,
and
27
July
1994
16:
evolution
(Cambridge University Press, London).
and two anonymous referees for their percep-
This
A
fertilization. Aust.
pro-
101-109.
(3rd ed.):