Determination of Sex

The essential feature of sexual reproduction is the productio~ of

male and female gametes._ This feature enables sexual reproduc!1on
to provide genetic variability. It is for !his reason th~t we bel~eve
that sexual reproduction has played an important role 1n evolution.
Among higher organisms, while in monoecious forms the male and
female reproductive organs are produced on the same individual, in
dioecious forms, they are borne on different individuals. In animals,
such a .difference in dioecious forms is accompanied with secondary
sexual characters. In plants, however, the male and female plants to
not differ except in floral characters. Moreover, dioecism in plants
is a rare phenomenon and most of the plants bear bisexual flowers
having male and female reproductive organs. However, in dioecious
forms, there is a specific . problem of finding out the factors which
are responsible for determining· whether a particular individual
would be a male or a female. This is actually the problem of
deter~ination of sex. In higher forms, it is observed that equal
number of male and female individuals are available in nature.
This ·would suggest that a very precise mechanism of sex
determination should be involved. Various mechanisms in different
forms of animals and plants are known and would be discussed in
this chapter.
CHROMOSOME THEORY OF SEX DETERMINATION
According to the chromosome theory of sex determination the
male and female individuals would differ in their chromos~me
constitution. There may be two types of chromosomes present
in such individuals : (i) autosomes, and (ii) sex-chromosomes.
In a diploid individual, there are 2n-2 autosoines and two
►

[ 261
DETERMINATION OF SEX
sex chromosomes are
sex-chromosomes. While in one sex, two
heteromorphic (XY),
homomorphic (XX), in the other sex these are
omosome mechanism
Table 16-1 shows different systems based on chr
of sex: determination.
mosomal determination of
Table 16-1. Different systems involved in chro
sex.
Gametes Zygotes
System
sperms eggs males I females
I
XO&' [ A+X(S0%) A+X(100%) 2A +xo 2A +xx
e.g. Protenor A+O (50 %)
XYo' [A+X!S0%) A+X (l00 %) 2A+ XY 2A +xx
e.g. Drosophila A+Y (50 %)
XY~
e.g. Birds
[A+ X
(100%)
A+X (50 %)
A+Y (50 %)
2A +xx 2A+ XY

XO~ A+X (50 %) 2A +xx 2A tXO

eA +X A+O (50 %)
e.g. Fornea 100 %)
on of sex became
-The role of sex chromosomes in the determinati
of sex linked genes
evident for the first time due to the discovery
ents of C. B. Bridges
discussed earlier in Chapter 14. The experim
ved that a diploid set
on the nondisjunction of X-chromosomes pro
2A+XX(~)' X 2A.+XY(O)
nondisjunction of ·XX I

.ga me tes

A+ X A+Y

2A+XXX 2A+xxv
A+XX superfemale female
(dies)

2A+X 2A+Y
A+O
male (dies~

Fig .. 16-1. Nondisjunction of X-chromosomes in

a female Droso hi/a
daughter. .. 'P
!~ad10g to th~ transfer of both X's to the !
262 ] CYTOLOGY, GENEtICS AND EVOLUTlO~

of autosomes with two X-chromos omes wilJ always give rise to a

female individual irrespective of whether the two X-chrotnos omes
came from the same parent or from different parents (Fig . 16-1).
The above example illustrates that the Y-chromo some does not
carry any sex determinin g factor. The presence of one or two
X-chromos omes is more important than the presence or absence of
Y-chromos ome.
BALANCE THEORY OF SEX DETERM INATION
The experimen ts discussed above, though have shown that the
Y-chromos ome is not important for the · determinat ion of sex, it
does not indicate whether X-chromos ome a!one determines the sex
or if the autosomes also play any r.ole in the mechanism . Individuals
could, however, be obtained, which had two X-chromos omes as in
the normal female, but were intersexes~. Each of these intersex:es had
an extra set of autosomes indicating that the autosomes play a
definite role in the determinat ion of sex. In this connection , Bridges'
experiment s on intersexes and supersexes are of special importance .
Bridges, as early as 1922, came .across certain Drosophila individuals
which were triploids and thus had three sets of chromosom es
(3A : 3X). These triploid individuals \yere normal females and were
crossed with diploid males (2A+XY). The results obtained from
such a cross are shown in Fig. 16-2. As is obvious, from such a
cross, normal diploid males, triploid females, intersexes, supermales
and superfemal es are obtained. The. chromosom e constitutio n of
these phenotypic ally different sexes has a definite bearing on the
mechanism of sex determinat ion.
[I] Triploid intersexe s in Droaophila and balance theory
The presence of triploid intersex:es in the experimen t conducted
by Bridges (Fig. 16-2) is a definite proof that the autosomes also
carry factors for sex determinat ion. These intersexes are sterile
individuals, which are intermedia te between male and female. The
results, where intersexes, supermales and superfemales were obtained,
were interpreted by Bridges in the form of Balance Theory of Sex
Determination. According to this theory, the ratio between the
number of X-chromos omes and the number of complete sets of
autosomes will determine the sex. The X-chromos ome is believed to
carry female tendency genes, while the autosomes carry the male
tendency genes. Both these sets of genes start functioning and there
hast~ be a balance between them for an individaal .to become male
or female. In one complete set of chromosom es (A+X); female
 -·, , I

Ir
I ,
lI

1JETERMINAt10N OF stx [ 26j

lI
3 A+ XX X (~)
triploid
X 2A+X Y(O)
diploid

gam etes
(oj 1'
'
A+X A+Y
I

I
3A+X XX 3A+X XY
2A + XX
triploid female triploid in1tersex

2 A.+ XX 2 A+ X Y I

I
I

A+X
diploid female diploJd ma le

3A-f. XX 3 A+ X Y
2A+X -
triploid intersex supermale i

2A+X XX 2A+X XY
·- . . ·- ..
A+X X
I

' . ·• . ' .. ..
superfemale. . diploid female
, .

-
Fig. 16-2. Result s obtain ed from a cross of a triploid (3A+X XX) ~ flY
and
a diploid (2A+X Y) i! fly in Drosophila.

Table. 16-2. Differe nt doses of X-chro mosom e· and autoso me sets and
their .effect on sex determ ination .
Plofd1 X Sets of . X/A
autosomes ratio
Jevel chromosomes
diploid 2 J ·50] super fem ale
3 .l ·33
tnploid . 4 3
haploid J.
diploid .
tnploid
tetrapl oid ·
tripJoid
tetrapJoid
2
3
4
2
3
u
3
4
1·00

0 67]
0·75
fem ale

interscx

diploid 1 2 0·50] male

tetrapl oid 2 4 O·SO
triploid l 3 0·33 supermale
 264 ] CYTOLOGY, GENETICS AND EVOLUTiO:N .

Table 16-3. A summ ary chart showing

different ratios of X-chromosornes and
autos ome sets (X/A ratio) controlling
sex in Drosophila.
X/ A ratio sex
>l·O super femal e
\ 1·0
l·0-0 ·5
female
intersex
0·5 male
<0·5 · super male

tendency .genes _are more . Conse-

quently if the ratio betw een X and
A is one, it will pe a female
individual. The different ratios
(X/ A) and t~~ir corre spon ding
phenotypes (sex) are show n in
,._--- v---- -' '----r ---J Table 16-2 and summ arise d in
female male
Table 16-3.
Fig. 16-3. A gynan drom orph As show n in Tabl~ 16-2, when
Drosophila fly showing female XI A ratio is 1·0, the individuat
and male parts. will be female ~nd when it is 0·5, it
would be male. Whe n this balance
is disturbed, the individual deviates from norm al male or
norm al
female. For instance, when the ratio (X/ A) falls between
I ·O and
0· 50, it would be inter sex; when below 0· 50, it would be supe
rmale
and when above l ·O it would· be superfemale (Table 16-3).
[II] Gync1ndromorphs in Drosophila
In Drosophila, occasionally flies are obta ined which ,hav e fema
le
char acter s in one part of the
body and male char acter s in the
J., ~x x rema ining parts (Fig. 16-3).
Such indiv idua ls are know n
- {ff?

/\' ~
~ c::::=:J---.~
as gyna ndro morp hs and are
believed to result due to the
loss of an . X-chromosome
"- - thli1 X chromosome
will be lost
in a parti cular cell durin g
development. If this event
Fig 16-4. The los1 of an X-chromosome happ ens durin
g first mitotic
during mitosis in a 2A+XX cell leading
to the deriv ation of two daugh ter cells
division of zygote, then one
one having 2A+X X and the other having of the two cells of the two-
2A+X . celled proe mbry o will have
266 ] . CYtot oGY, GENEtlCS ANO EVOLUTION

that female . Europeans were designated as FwM w and male

Europeans as MwMw. This means that Fw chrom osom e of weak
race can overcome male ·determining factors in M w to produ ce a
normal fe1i1ale. Similarly in strong Japanese race, females are Fs Ms
and males are Ms Ms. Crosses between males and females of sam_e
race 0oive males and females in equal propo rtions . However, tf
weak and strong races were intercrossed, .different results were
obtained in reciprocal crosses (Figs. 16-5, 16-6).
It is obvious that in a cross between weak Europ ean female
(Fw Mw) and strong Japanese male (Ms Ms), intersexes (Fw Ms)
are obtained, because Fw can not overcome the male factors in Ms.
In the reciprocal cross between strong Japanese female (Fs Ms) and
weak European male (Mw Mw), no intersexes are obtained.
SEX DETE RMIN ATIO N IN PLANTS AND MAN
In plants, as also in case of · human beings, the sex is mainly
controlled by the Y chromosome. Consequently, irrespective of the
number of X-chromosomes and the number of sets of autosomes, if
Y-chromosome is present, the individual would be male while if
Y-chromosome is absent, it would be female. Evidences for the
control of sex by the Y chromosome will be presented separately for
plants and for human beings (Homo -sapiens).
(I] Plants
The mechanism of sex determination has been studied in a large
number of plant materials. A review of the mechanism of sex
determination known in flowering plants was presented by
M. Westergaard. A detailed account of sex-determination in plants is
also availab]e in C.R. Burnham's book "Discussions in Cytogenetics".
While a number of dioecious species have so far been examined,
detailed study was undertaken in Coccinia recently in India and in
Me/andrium e]sewhere. Therefore a somewhat detailed account of
sex-determination in Coccinia and Melandrium will be first presented
and a summary of other plant species examined in this manner will
be presented separately,
1. Coccinfa indica and Melandrium alb"m ( Lychnis) : The
mechanism of sex determination in Coccinia indica a member of
family Cucurbitacea~ has been studied in s~me detail by
Prof. R.:. ~oy ~nd ~ts c~-workers at Patna University. They studied
the sex m d1plo1d, triploJd and tetraploid plants with and without
Y chromosome and observed that irrespectiv~ of the number of
 ~
I

DETERMINATION OF SEX [ 269

3. Other dioecious plants. In above examples of Coccinia indica

and Melandrium album, male sex was heterogam etic (~XX;(! XY).
Other examples having different kinds of sex chromoso me
constituti ons are listed in Table 16-6.
[II] Homo sapiens
1. Y-chromosome in sex determination. (Turner's and
Kliuefelter's syndromes). In Homo sapiens (human beings), as in
,
J

7
' Melandrium, Y-chrom osome controls the sex. Conclusive evidence
in this connectio n came from certain abnorma l individuals (caHed
., syndromes). Turner's syndrom es (XO) named after the person who
discovered them are sterile female individuals having certain abnor-
[,
td
malities. Similarly , Klinefelt er's syndrome s (XXY) named after
of Klinefelter are males, despite the presence of two X-chromo somes ..
l0-
A Drosophila fly with 2A+XXY , which is exactly like Klinefelter''s
mt Table 16-6. Chromoso me constitutions of two sexes in some dioeciou pfaBt
css species.
ing Examples
Mechanism

!lg 1. Male heterogam etic Humulus lupulus (hops)

I
Melandrium album, M. rubrum,
Uld __...
(~ XX; a
XY)
. Rumex angiocarpus
we iess
well { Populus spp., Salix, Smilax, and
t on established Cannabis
2. Male heterogam etic Vallisneria spiralis
ous, a
(~ XX; XO)
Dioscorea sinuata
Phoradendron .flavescens
van. 3. Male heterogam etic
(see (~ with an extra chromosom e) P. villosum
, for 4. Female heterogam etic Frogaria elatior and
other Fragaria species
s, 14 a
(~ XY; XX)
R11mex acetosa
>mo-
),
,. Compound chromosomes
Hum11/us Japonic11s
ially, (e.g. ~ XX; i
Y1XY2)
6. No ,ex chromosomcw Splnacla ot, racea
ise·to Rlbes alplnum
(perhaps gene contro11ed)
omo- Y/11.t ctn,rtta
(11*) V. r11p1Jtrls
V. vlnlfera
ith 11 Car/ca papaya
ations ibpllrtJg U.f ,1ffidnal/s
tmetes Bryon/a dio /cc•
10n in J,'/.tc11m Jbcheri
7. No 1ex chromoaom c,
in male
(translocati on hcteroiysoaity)

1,
 <

270 ) CYTOLOGY, GENETICS AND EVOLUT

ION
syndrome of human beings in chromoso
me constitution, is a normal
female individual. This obviously sugges
ts that in human beings the
sex determining mechanism is essentially
like that in plants (Coccinia
and Melandrium) and not like that in Dro
sophila.
2. Rare XY females (due to deletion)
and XX males (due to
translocation) in humans. In 1986, res
ults have been published,
which showed that XY females may
be produced which have a
deletion in the region lA on the sho
rt arm of Y chromosome.
Similarly XX males were discovered, wh
ich had the segment lA of
the stort arm of Y chromosome translo
cated to one of the X chro-
mosomes. This led to the conclusion
that a segment on the short
arm of Y chromosome determines malen
ess and not the ge.ne for
H- Y antigen (as was earlier believed)
located either near ·the
centromere or on the long arm.
SINGLE GENE CO NT RO L OF SE
X
In the examples discussed in the previous
sections of this chapter
the sex was determined by- individual sex
chromosomes (X or Y) or
by a complete set of autosomes. rt is
also known that, in s11ch
cases, whole chromosome having large num
ber of genes ratheI than
single gene controlled the ·sex. Howeve_
r, there are other instances
where individual single genes-we~e found
to be responsible for the
determination or expression of sex. · Three
such cases will be briefly
described in this section.
[I] Sex de ter mi na tio n in Asparagus
Asparagus is a dioecious form. Howev
er, rarely the female
flowers bear rudimentary anthers and
the male flowers bear rudi-
mentary pistils. Thus, the rare male flow
ers having poorly developed
pistils may set seeds. In one such cas
e when the seeds obtained
Rare male plant

I••lf•d
from a rare male flower were raised
(Pp) into plants, the male and female
plants, were found to be present in
I I 3 : l ratio. When the male plants
PP Pp PP raised thus were used to pollinate
25
l ( ¾> (50¾) ' 25
( ¾) the female flowers on female plants,
0~ (1;•.1,, r, (25'-')
'' • '' only two-third of them showed
Fig. 16-8. Segresation for se,c segregation indicating that the sex
in seed obtained from a rare is controlled by a single gene. In
bisexual flower in Asparagus this case, maleness should be domi-
showing monogenic control. nant over femaleness and the