Genetica (1967) 38:388-397

CYTOMORPHOLOGICAL STUDIES IN THE FAMILY

SOLANACEAE FROM WEST PAKISTAN

S. R. t3AQUAR
Cytogenetics section, Central Laboratories
P.C.S.I.R., Karachi-32, Pakistan
(Received July 6, z967)

A project on the chromosome survey of the flora of West Pakistan is in

progress and chromosome counts of a large nunlber of species from this
region have been reported earlier (BAQUAR et al. 1965, 1966, 1967). The
present paper is a part of this project. Morphological and cytological be-
haviour of 29 species distributed over 12 genera of the family Solanaceae
have been investigated for the first time from West Pakistan. The present
investigation shows that the cytological behaviour of the plants of this re-
gion is almost similar to those of the other parts of the world. ]Besides the
confirmation of the earlier findings, this report includes four new records
namely: Solanum albicaule KOCSCHY n = 12; Solarium gracilipes DECNE
n = 12; Withania coagulans DUN~L n -- 24; Datura metel L. n ~ 12,

Introduction
TjIO (1948) r e m a r k e d t h a t the " d e v e l o p m e n t in p l a n t c y t o l o g y
d u r i n g t h e last half c e n t u r y has been m a i n l y m a d e on m a t e r i a l from
t e m p e r a t e or arctic zones, while t r o p i c a l p l a n t s h a v e been r a t h e r
s t e p m o t h e r l y t r e a t e d " . H e b e l i e v e d t h a t m a n y cytological p h e n o m e n a
m i g h t receive a different outlook after this gap in our c y t o l o g i c a l
knowledge is bridged. U n f o r t u n a t e l y such i n f o r m a t i o n on t h e flora
of W e s t P a k i s t a n is y e t negligible. U n d e r a p r o j e c t on t h e c y t o m o r -
phological s u r v e y of t h e flora of W e s t P a k i s t a n , c h r o m o s o m e n u m b e r s
of a large n u m b e r of species h a v e been r e p o r t e d earlier (BAQUAR et al.
1965-i967). The p r e s e n t w o r k is a p a r t of t h a t p r o j e c t in w h i c h
chromosome counts of 29 species of t h e f a m i l y Solanaceae, including
four new records, h a v e been r e p o r t e d .
T h e i m p o r t a n c e of t h e Solanaceae, because of t h e i r economic a n d
m e d i c i n a l value, is well known. T h e y include m a n y food p l a n t s
(potato, t o m a t o , p e p p e r etc.) a n d m e d i c i n a l p l a n t s (henbane, b e l l a d o n a
n i g h t shade etc.), a p a r t from tobacco a n d m a n y o r n a m e n t a l plants.
Out of 85 genera a n d 2200 species (PORTER 1959), d i s t r i b u t e d t h r o u g h -
out t r o p i c a l a n d t e m p e r a t e regions, it is r e p r e s e n t e d b y o n l y 12 g e n e r a
 CYTOLOGY OF S O L A N A C E A E FROM PAKISTAN 389

and about 35 species in West Pakistan. All the earlier reports on the
chromosome counts of the members of Solanaceae are from non-
Pakistani material except those reported earlier by tile present author.
In the present report, the chromosome numbers and references
already given in DARLINGTON & WYLIE (1955) and L6vE & LOVE
( I961) have been omitted.

Materials and Methods

Chromosome numbers were determined mainly at meiotic meta-

phase of the pollen mother cells using the usual Carnoy fixative
(alcohol-chloroform-acetic acid, 6 : 3 : 1) and the acetocarmine
squash technique, with the exception of Hyoscyamus niger L., Datura
stramonium L. and Datura metaloides DUNAL. In these three cases
counts were made from root-tips, which were obtained from seeds
grown on moist filter paper in petri dishes at room temperature
(25~176 Root-tips, about two cm long, were excised and pretreated
with 8-oxyquinolin for three hours before fixation. Staining and
squashing were carried out using the Feulgen aceto-carmine squash
method.
The materials were collected from plants growing in their natural
environment in the field, from the localities mentioned against
respective species (Table 1). Apart from the materials collected by
the author himself, seeds and fixed buds were received from various
collegues to whom the author gratefully wishes to acknowledge.
Localities: G = Gharo (in Indus delta), K ---- Karachi,
M = Murree (in Punjab), P = Peshawar (in NWFP),
PI = Pipri (near Karachi), T = T h a t t a (in Indus delta),
RP = Rawalpindi (in Punjab).

Cytological Observations
Solarium
Out of the 11 species of Sola~um investigated (Table I), eight are
indigenous while the remaining three namely, S. melongena L., S.
pseudocapsicum L. and S. tuberosum Lo are cultivated. All the species
show a uniform gametic number of I2, except S. tuberosum in which
it was found to be 24. The chromosome counts for S. albicaule KOT-
SC~IY and S. gracil@es DECNE. are new records while the rest are in
390 s.R. BAQUAR

TABLE 1
CHROMOSOME NUMBERS (n, 2n) AND PLOIDY (X) IN Solac4cl,c6ae FROM THE PRESENT
INVESTIGATION COMPARED WITH PREVIOUS REPORTS

Species Present Investigation Previous reports

Locality n x n 2n A u t h o r i t y *)
Solanum
nigrum L K, R P 12 4 12 ~X~ANDA (1962), CHUANG et al.
1963), TANDON • RAO (1964),
BORGMAN (1964),
BA~UAR et al. (1965)
20 RAI (1959)
24 TANDON & RAO(1964)
36 MULLIGAN 1961),
MASUBUCHI (1961),
TANDON & RAO (1964),
TURALA (~ URBAN.-WoRY in
SKALINSKA et. al. (1964)
72 GADELLA & KLIPHUIS (1967)
surattense
Burm.f. K, T, R P 12 4 12 24 BEZ. & BEZ. (963),
BAQUAR et. aI. (1965)
incanum L. K, P I 12 4 24 MIa~GE (1962)
albicauble
Kotschy K 12 4
gracilipes
Decne. K 12 4
dulcamaraL. R P , M U 12 4 24 GADELLA ~: t~LIPHUIS (1966)
indicum L. RP 12 4 12 24 BEZ. & BEZ. (1963) CHENN. & K. (1965)
verbasci-
]olium L. M U 12 4 12 24 BEZ. & BEZ. (1963)
melongenaL. IZ 12 4 12 CAPINPIN et al. (1963) CHENN. • K. (1965)
24 KAPOOR & TA~DON (1964),
(2 n x 2) C~tENN. & K. (1965)
pseudo-
capsicum L. R P 12 4 12 24 BEZ. &BEz. (1963) KAWANO (1965)
tubero-
sum L. K 24 8 48 GILLES (1960) (YEN & P 1965).
96 WANGENHEIM (1954)
Withania
somnifera
L. DUNAL K, T 24 8 24 GOTTSCHALK (1954).
MOHAN RAM (I964).
BAQUAR et al. (1965)
48 MI]kGE (1960)
 CYTOLOGY OF SOLANACEAE F R O M P A K I S T A N 391

T A B L E 1 (continued)

Species Present Investigation Previous reports

Locality n x n 2n A u t h o r i t y *)

coagulans
DUNAL K 24 8 --

Physalis
minima L. K, R P 24 8 24 BAQUAR et al. (1965)
peruvianaL. R P 24 8 12 GOTTSCHALK (1954)
24 SHIBATA (1962), HEISER (1963)

Lycium
eu~'o-
paeum L. K, T, G 12 4 12 MALIK (1960),
BAQIJAE et al. (1965)
18 6 18 BAQUAR et al. (1965)

Hyoscyamus
niger L. P 34 2 34 See LOvE & LOv]~ (1961)
2n
ADal'g,ga,
metel L. K 12 4 m

stramo-
nium L. P 24 4 24 See LOvE & LOvE (1961)
innoxia 2 n
MILL. RP, MU 12 4 24 SOBTI & SINGH (1961)
metaloides
DUNAL RP 12
P 24 4 24 See LOvE & LOvE (1961)
2n
A tropa
acuminata
I~OYLE e x
LINDL. RP 36 12 36 MEHRA • SOBTI (1954)

Nicotiana
plumbagini-
/olia Vlv. R P ca 21 20 GOODSPEED (1945)
tabacum L. RP 24 8 48 See LOvE & LOVE (1961)
rustica RP 24 8 48 See LOvE & LOVE (1961)

~est~'um
~OC[,~td/-
hum L. K 48 2 14,16 SI-:ARMA & S~IARM* (1958)
392 S. R. BAQUAR

TABLE 1 (continued)

Species Present Investigation Previous reports

Locality n x n 2n Authority *)
Petunia
Petunia
hybrida K 7 2 7 GOTTSCHALK (1954)
14 TAKEHISA(1961, 1964)
16 TAKEHISA (1961)
21-28 TAKEHISA(1961)
28 BREUER(1961),
1V][ORRISON (Y~ RAJHATHY (1960)
Capsicum
annuumL. K 12 4 12 BAQUAR et al. (1965),
24 OHTA (1962)

Lycopersicon
esculentum
MILL. K 12 4 24 RICK (1960), DIERS (1961),
MASUBUCHI (1961),
LIMA-DEPARIA& SARV. (1962),
48 MORRISON & RAJHATHV (1960)

*) Authors quoted in DARLINGTON& WYLIE (I965) and LOVE & LOvE (1961)
have not been included in the list.

a c c o r d a n c e w i t h t h e earlier reports. Meiosis was n o r m a l a n d the

chromosomes were well defined ill all t h e species. DARLINGTON &
WYLIE (1955) r e c o r d 12 a n d 23, whereas LOvE & LOve ( 196 I) consider
6 as basic n u m b e r for this genus; x =- 23 has been r e p o r t e d in o n l y
two species viz. S. aviculare (2n = 46) a n d S. laciniatum (2n ~ 92)
b y BAYLIS (1954).

Withania
This genus is r e p r e s e n t e d b y only two species in P a k i s t a n n a m e l y
W. somni/era (L.) DUNAL a n d W. coagulans DUNAL. T h e former is t h e
p r e d o m i n a n t l y occurring species, a t h i n l y woolly shrub, of v a r i a b l e
shape, h a v i n g ovate-oblong, t h i n green leaves, h e r m a p h r o d i t e flowers
a n d berries loosely enclosed b y t h e calyx. The l a t t e r is a c o m p a r a t i v e l y
rare u n d e r s h r u b a n d is distinguished from t h e former ill h a v i n g
oblanceolate-oblong, thick, whitish leaves, dioeeous flowers a n d w i t h
berries t i g h t l y enclosed b y t h e calyx. B o t h t h e species h a d n = 24.
The counts for W . coagulans are a new record as it does n o t a p p e a r to
 CYTOLOGY OF SOLANACEAE FROM PAKISTAN 393

have been i n v e s t i g a t e d as yet. Meiotic b e h a v i o u r of W. somni/era was

v e r y a n o m a l o u s showing all sorts of irregularities such as laggards,
bridges a n d precocious units.

Physalis
P: minima L. a n d P. fleruviana L. are the only two representatives
of this genus t h a t are f o u n d in West P a k i s t a n . The former is a weed.
Physalis peruviana is a c u l t i v a t e d species. The chromosome n m n b e r s
observed (n = 24) agree with earlier reports.

Lycium
Lycium europaeum L. is a s p i n y shrub found a b u n d a n t l y in s o u t h e r n
West P a k i s t a n besides its occurrance in the n o t h e r n zone. I t displays
a great cytomorphological diversity.
There is some confusion about the validity of the name of this taxon. HOOKER
(1883) described two closely resembling species, L. europaeum and L. barbarum
L. from "~V.Asia and expressed his doubt "whether either of these two ought to
be separated specificaIly". Cook (1908) determined the plants of Karachi and
Sind as L. barbarum L. instead of L. eu~1opaeum and said it seems hardly separ-
able from L. europae~m. Another name, L. edgeworthii DUNAL has also been,
at times, given to this taxon. JAFRI (1966), prefered L. europae~tm and described
L. bc~rbar as a synonym, for the specimens of the Karachi flora. ]~AQIdARet
al. (1965) reported the chromosome number (~z = 12 and n = 18) of this taxon
under the name L. barbc~r~nr for plants from the Indus delta. This is main~cained
for this report.

Hyoscyamus
Out of the two species of Hyoscyamus reported from West P a k i s t a n i
only one, H. niger L. has been investigated, while the other, H. muticus
L., has n o t yet been collected b y the author. The chromosome n u m b e r of
H. nigerwas d e t e r m i n e d from the m i t o t i c m e t a p h a s e in which 34 chromo-
somes w ere counted, confirming earlier reports (L6vE & L6vE, 1961).

Datura
Datura is an a n n u a l or b i e n n i a l tall herb with large t u b u l a r flowers
a n d prickly capsular fruits a n d is f o u n d c o m m o n l y growing near
waste places. There are four species of this genus in West P a k i s t a n :
D. innoxia MILL., D. stramonium L., D. metaloides DUNAL a n d D.
mete! L. The last m e n t i o n e d species is restricted to the s o u t h e r n region
while the others are d i s t r i b u t e d in the n o r t h e r n part of W. P a k i s t a n .
A gametic n u m b e r n = 12 was recorded in D. metel a n d D. innoxia
394 s.R. BAQUAR

I 2 3 4
Figure 1. S. albica~le KOTSCi~Yn = 12 (iV~II).
Figure 2. S. gracilipes DEc•E n = 12 (1ViII).
Figure 3. Withania coagulans DUNAL n 24 (IV[I).
=

Figure 4. Datura mete[ L. n ~ 12 (IV[I).

a n d 2n = 24 in D. stramonium a n d D. metaloides, which is in accord-
ance w i t h the previous findings. T h e count for D. metel is a new record.

Atropa
Atropa acuminata ROYLE ex LII~DL. A t diakinesis 36 b i v a l e n t s were
observed, in a c c o r d a n c e w i t h the previous r e p o r t of MEHRA & SOBTI
(1954).
Nicotiana
N . tabacum L. a n d N . rustica L. are c u l t i v a t e d for t o b a c c o in
P a k i s t a n . The chromosome n u m b e r s have been r e p o r t e d r e p e a t e d l y
(Table 1), a n d t h e p r e s e n t result of n = 24 for b o t h species, c o u n t e d
a t M I a n d M I I confirm t h e previous findings. I n N . plumbaginifolia,
t h e only species of N. t h a t has e s t a b l i s h e d itself in t h e wild s t a t e in
P a k i s t a n , t h o u g h meiosis was normal, t h e chromosomes were difficult
to count. A g a m e t i c n u m b e r of a p p r o x i m a t e l y 12 has been d e t e r m i n e d ,
which is different to t h e o n l y previous r e p o r t (GooDSPEED, 1945) of
2n = 20. I t is certain t h a t the n u m b e r is n o t less t h a n 10 nor m o r e
t h a n 12.
Cestrum
C. nocturnum L. A g a m e t i c n u m b e r n ---- 8 was c o u n t e d at M I in
t h e P.M.C., which is in line w i t h t h e earlier r e p o r t of SHARMA
SHARMA (1958) who r e c o r d e d 2n = 14 a n d 16.
Petunia
Petunia hybrida. T h e record of n ~ 7 is a c o n f i r m a t i o n of previous
findings, including 2n ~ 14, 16, 28 (Table 1).
 CYTOLOGY OF S O L A N A C E A E FROM PAKISTAN 395

Capsicum
Capsicum annuum L. The present report of n = 12 confirms earlier
reports including 2n = 24, 36, 48 (Table 1).

L yc@ersicon
L. esculenlum MILL. The record of n = 12 is a confirmation of
previous records (Table I).

BASIC NUMBER

The question of the evolution of chromosome numbers in the family

Solanaceae poses great inconvenience due to the presence of m a n y
unrelated somatic numbers as 12, 14, 16, 18, 20, 21, 22, 24, 25, 28, 31,
34, 35, 36, 38, 40, 42, 44, 46, 48, 50, 60, 64, 72, 92, 96, 144. This in-
consistancy is restricted to a few species, especially the cultivated
ones, and not all the genera. Out of 14 genera described from central
and northwestern Europe I0 are based on x = 6 (L6VE & L6VE 1961),
DARLINGTON & WYLIE (1955) mention n = 12 for these genera.
Nicotiana, cultivated throughout the world and under constant
experimental strain since time immemorial, displays the greatest
inconsistancy (18, 20, 24, 31, 36, 38, 44, 48, 64, 72, 96). In the presence
of the experimental number of 2n = 12 in the genera Capsicum
(CHRISTENSEN • ]3AMFORD 1943), Datura (SATINA et al. 1941) and
Lycopersicon (RICK 1945), it is presumed that 6 is the original basic
number in the family from which 7, 8, 9, 10, 11 and 17 have evolved
secondarily, during the course of time, b y addition or reduction of
chromosomes in the diploid and tetraploid.

The author wishes to acknowledge Dr. A. IZAMAL, Director, Central

Laboratories for providing facilities and Mr AEAQ HUSAIN for tech-
nical assistance during the progress of this work.

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