06_chapter 2

CHAPTER
REVIEW
Although
com paratively
received
has
in
selection.
The
simply
is
being
of
it
is
entirety,
present
as
The
an
of
attention
Since
its
com pleteness.
that
induction
extensive
accum ulated.
taken
LITERATURE
a nsu branch
literature
at
OF
2
m utations
cytogenetics,
and
a vast
not
possible
it
review ,
literature
illu stration
conducted
by
and
is
polyploidy
it
has
quantity
to
necessary
of
does
not
mentioned
here
is
various
the
literatu
nature
this
make
therefore,
of
already
review
to
of
investigators.
is
a
aim
to
work
be
13
2.1,
INDUCTION
OF MUTATIONS
Cereals
S a r i c et
(1961)
stud ied
the e f f e c t of
gamma
r a y s on wheat seeds with respe ct
to the morpho- p h y s i o l o g i ­
cal
and found that
c h a r a c t e r i s t i c s of
investigated
the
germination as well as he ig ht
s e e d l i n g s d ec rea se d with i n c r e a s e
v a r i e t i e s te ste d
towards the
Pa lenz ona ( 1 9 6 1 )
the
"reversal
seedling
observed
phenomenon**,
m ois tu re level
B or o j e v ic
of
Reaction of
in uheat s e e d l i n g s
a sharp
of
achieved
was
se ed s .
and B or oj ev ic
(1969)
studied
stabilization
in i r r a d i a t e d p o p u la t io n of
The r e s u l t s showed that
r ea chi ng
s t a b i l i z a t i o n of
the Mg
the mean v a l u e s of the control
constancy in l a t e r
Kivi
and a
s pe c ie s and on the
ch a ra ct er s i n i r r a d i a t e d p o p u l a t i o n began i n
generation,
that
de pr es s io n of
recovery at very high d os es ,
induced g en et ic v a r i a b i l i t y
v u l q a r e wheat.
and weight
same dose was d i f f e r e n t .
that i s ,
depe nd ent on the p lo id y leve l
the
in d o s e .
growth dependent on the r a d i a t i o n dose
subsequent p a r t i a l
of
some of the
doses were stimulant and others i n h i b i t o r y .
The percentage of
of
s e e d l in g s
and
g e n e r a t io n s *
and Ramm-Schmidt ( 1 9 6 9 )
obtain ed
sp routing
r e s i s t a n t mutants from Co60 i r r a d i a t e d po pu la t io n of
Finnish
spring wheat variet y
Kapoor and N a t ar aj an
Ru se.
(1971)
h e x a p l o i d bread uheat ( Triticum
ex p o s u r e s of
gamma r a y s .
grew s ix v a r i e t i e s of
aestivian )
under chronic
Among v a r i e t i e s t e s t e d ,
three
14
varieties
(N .P ,
797,
798 and 8 3 6 )
under i d e n t i c a l
responded d i f f e r e n t l y with r e f ere n ce
plant
height,
seed f e r t i l i t y ,
conditions
to ch a ra c te rs l i k e
morphological
a b n o r m a li t ie s
and v i a b l e m u t a t i o n s ,
Peix ot o
varieties
Gomes ( 1 9 7 2 )
irradiated
with the o b j e c t i v e o f
w ith out a l t e r i n g
seeds of
three wheat
sh ortening t h e i r
straw
other important c h a r a c t e r i s t i c s ®
sh o r t strau mutants were o b t a i n e d ,
improved r e s i s t a n c e to Pu c c in ia
No
but mutants w ith
cram in is t r i t i c i
were
observed,
Singh ejb a l .
(1972)
observed in mutation
on bread wheat ( Trlticum aestlvum) .
p h y s i c a l mutagens,
in
using chemical
as well
that the treatments which had r es ul t e d
high er s t e r i l i t y
in
gave hi gh er mutation percentage
o f phenotypic d e t e c t a b l e types i r r e s p e c t i v e of
dosage e f f e c t ,
studies
varietal
treatment,
response or geno-type treatment
interaction*
Shakvarnikov jat a l .
v a r i e t i e s of T ,
o f physical
mutations*
radiation
(1976)
s u b j e c te d d i f f e r e n t
aestlvum and T* durum to a l a r g e
and chemical mutagens for in d u c ti o n
Different varieties
viz*,
h e a vi e r g r a i n s ,
better
baking q u a l i t i e s ,
Valeva ( 1 9 7 6 )
higher
to
some good economic
g l ut i n c o n t e n t ,
were scored,
studied induced mutations i n wheat
at v a r i o u s pl o id y l e v e l s ,
h e x a p l o i d with gamma
etc*
of
responded d i f f e r e n t l y
treatments* Mutants p o s s e s s i n g
traits,
number
that i s ,
diploid,
tetraploid
rays and e t hyl en e i m i n e .
and
Presenc e
15
of
varietal
response was found dependent on ge notypic
constitution
results it
ploidy
in a d d i t i o n to p lo id y l e v e l s .
was concluded that the s p e c i es f r o *
groups,
resistance
From the
on the whole,
higher
showed a greater degree of
towards mutagenic treatments with regard to
parame te rs s t u d i e d . Yet they might show higher m u t a b i l i t y
if
the genotypes s el ec te d are c u l t i v a t e d v a r i e t i e s *
Puzakova _et jjJ,,. ( 1 9 7 8 )
s tu d i e d e f f e c t of
i r r a d i a t i o n on wi nt er hard ine ss o f
wheat.
The doses which i n h i b i t e d
*ulyanovka*
amino a c i d s ,
wint er
growth promoted
s im ul ta ne o us ly the accumulation of reducing
free
gamma
sugars and
and in the autumn made the p l a n t s more
hard y fo r s u r v i v a l .
Sharma and
with d iffe ren t
and
hydraz ine
Gupta ( 1 9 7 8 )
tr ea te d
doses of gamraa r a y s ,
(0.50$)
for d i f f e r e n t
seeds o f
T.
durum
ethyl methane'"'sulphonate
du rations to
demonstrate t h e i r cytogenetic e f f e c t s .
LD 50 was observed
to
l i e between 6 hr and 12 hr treatment for EI*IS whereas
it
was below 6 hr treatment for h y d r a z i n e .
Structural
a b e r r a t i o n s such as t r a n s l o c a t i o n s were observed only in
10
Kr and 30 Kr gamma ray tre atm en ts.
As ea rly
as 1 9 2 8 ,
S ta dl er
reported
artificial
p r o d u c t i o n of mut ations in b a r l e y by the use o f X and
gamma r a y s .
Almost a l l
and of no p r a c t i c a l
the mutations induced were l e t h a l
value.
16
Gustafsson
useful
(1947)
induced mutations in ba r l ey
as h e ig ht of s tr aw ,
straw,
chemical
protein
con ten t,
In bar ley
^2
observed the occurrence of
earliness
properties,
1,000
Gaul
for such c h a ra c t e rs
and l a t e n e s s ,
st r e ng th of
brewing c h a r a c t e r i s t i c s ,
grain weig ht and t i l l e r i n g
(1961)
observed
ca pa ci t y*
that the frequency of
chloro phy ll mut ations decreased s i g n i f i c a n t l y
in l a t e r
formed t i l l e r s *
Heslot e t & 1 .
substances like
in barley
Gaul
(1961)
concluded
that radiomimetic
EMS were more e f f e c t i v e
than i o n i z i n g
(1965),
radiations*
a f t e r measuring
chlorophyll mutations,
in in d u c i n g mutations
observed
mutat ion . frequency was l i n e a r
the frequency o f
that the i n c r e a s e in
with i nc r ea se
in
the dose*
EMS appeared to be a more powerful mutagen than X-rays*
U a l t h e r (1 9 6 9 )
mu t at io ns in the
considered the number of ch lo rop hyl l
ge ner ation as the b a s i s f o r
e f f i c i e n c y of X-rays and thermal
dormant seeds of b a r l e y .
After
calculating
neutron i r r a d i a t i o n
on
the s t u d i e s he proposed
f a c t o r of e f f e c t i v e n e s s which could give inf o rm a ti o n
a
on
the number of mutations per 100 i r r a d i a t e d seeds'*
Gaul e t ,al.o(l969)
observed that micro- m ut ati on s
are more important for d ir ec t use in pl an t b r e e d i n g
macro mutations*
to
They showed that i t was r e l a t i v e l y easy
s e l e c t micro-mutations having
hiqher
than the mother va rie ty
y i e l d p o t e n t ia l
than
a yield potential
and i t was expected
can be fur the r r a i s e d in s ec ond ,
10$
that the
t h i rd
and
17
later
was
cyc les o f mutagenic treatment by s e l e c t i o n .
suggested that
newly
for the continuous improvement of
r ele ase d v a r i e t i e s ,
the micro-mutation
It
particularly
the top v a r i e t i e s ,
techniques should be used and combined
w it h maintenance b r e e d i n g ,
Sharma and Bansal
(1971)
showed that mutation
f r e q u e n c y in b a r l e y was higher when i t s EMS t r e a t e d
were d r i e d back in comparison to those sown in
seeds
high
moisture conditions*
Doll
Cvar,
in
(1972),
Carlsberg
II)
after treating
the seeds o f ba r l ey
with EFISf induced a l a rg e v a r i a t i o n
dry matter y i e l d but no mutant was observed with a
s i g n i f i c a n t l y h ig h e r y i e l d of p r o t e i n s or non-protein
m att er per u n i t
lines
area.
dry
The non-protein y i e l d o f mutant
was found to be reduced more than the p r o t e i n y i e l d
dU8 to which an i n c r e a s e in percentage of p r o t e i n in most
of
the mutant l i n e s was n o t i c e d ,
Gustafsson et .al.
(1972)
were able to i n c r e a s e
dormancy pe r io d s in bar ley v a r ,
short
K r i s t i n a by mutagenic
tr e a t m e n t s .
Hansel ejb
(1972)
t r e a t e d seeds o f a cross-bred
s t r a i n of spring b arl ey with thermal neutrons and EMS?.*
Fro m
1^2
Ng p o s i t i v e
selection
for yejlld and kernel
w i dt h was c a r r i e d out under va ry i ng s e l e c t i o n p r e s s u r e s .
R e s u l t s with s e l e c t i o n
better
for y i e l d
a f t e r neutron i r r a d i a t i o n
in early g e n e r a t i o n s were
than a f t e r EMS treatment
b ut in l a t e r g e n e r a t i o n s the b e s t l i n e s of both treatments
gave
s i m ila r r e s u l t s .
18
Ooshua ejt *|1.
( l 9 7 4 #"a)
ob se rve d that combined
tre atment of b a r le y u/ith f a s t neutrons and d i e t h y l
sulphate
r e s u l t e d in s y n e r g i s t i c e f f e c t s on both anaphase chromosome
fragments and bridges#* Storage of
seed3 a f t e r neutron
i r r a d i a t i o n had no s i g n i f i c a n t e f f e c t on frequency of
frag ments and b r i d g e s .
3oshua jjjt a l .
of
(1974b)
f u r t h e r stud ied
the e f f e c t s
f a s t neutrons and gamma rays on s e ed li n g h e i g h t and
chromosome a b e r r a t i o n s in b a r l e y .
In response
to the
tre a tme nt s a s y n e r g i s t i c e f f e c t on s e e dl in g h e i g h t was
noted.
Further,
anaphase b rid ges
the presence of
chromosome fragments and
showed an a d d i t i v e e f f e c t wi th combined
treatments.
Thakare jst 1 g L . ( l 9 7 4 ) , a f t e r combined treatment of
barley
seeds with thermal neutrons and
a alight s y n e r g i s t i c e f f e c t in
'• dES ,
ob ta ine d
s e e d l i n g heig ht at lower
doses and p r o t e c t i v e a f f e c t at h ig h e r dos es,
wh il e
combined
treatment of gamma rays and dES gave s y n e r g i s t i c e f f e c t
on
s e e d l i n g h e ig h t at a l l the dos es t e s t e d .
Sethi ( 1 9 7 5 )
tr ea te d the
v a r i e t y o f ba r le y with EMS,
dormant seeds of C 164
gamma r a y s ,
P
32
and S
3S
,
147
v i a b l e mutant types were scored in H^ and confirmed in
generation*
These in clu de d D w a r f ,
Very early.
Early,
Late
leaf
Dark
green l e a f ,
Erectoides,
Profuse t i l l e r i n g ,
Erect l e a f ,
L i g u l e l e s s cum - aur icl ele ss,
Thick g re en ,
straw,
Synchronous t i l l e r i n g 1*
s e ne s ce nc e, Long broad l e a f ,
culm , Male s t e r i l e ,
Stiff
Double
Thick
> floret,
stiff
Brittle
19
awn ,
Multinodosum-cum-monopodial branching^ Many noded
dwarf,
Thick calm- dwarf and Long peduncled d w a r f .
M ut a nt s obtained
were m e i o t i c a l l y normal and f u l l y
fertile.
I n f l u e n c e o f DMSO (dimethyl
su lp ho x id e)
on the
m u t a g e n i c i t y of CMS in barley was studied by Kh al a tk ar
( 1 9 7 6 ) , Dry and
soaked
seeds were t rea ted with EMS alone
and in combination with DM50.
indicated
and M2 g e n e r at io ns
that DMSO was not a good c a r r i e r for
EMS + DMSO treatments a s l i g h t
EMS,
With
decrease i n mutation
fr e q u e n c y in M^ was obs erved.
L i ejb a l .
mutant l i n e s i n
(1961)
utilized
X-radiation to produce
10 v a r i e t i e s of r i c e b el ong ing
to both
j a p o n i c a and i n d i c a t y p e s 0 From s i x of the 10- X- irr ad ia te d
v a r i e t i e s 37 promising l i n e s were
of
derived.
These con sis te d
20 high y i e l d i n g m or ph ol ogi cal ly and p h y s i o l o g i c a l l y
normal l i n e s ,
but
S short-culmed,
3 e r e c t o i d ( s h o r t culmed
with markedly high s t i f f n e s s of
4 multibranched
and 2 ea rly
the culro)#
types which all e x c e l l e d
their
mother
v a r i e t i e s in y i e l d .
In one of the X-ray tre ated
indica
va ri et y three non-shattering mutant l i n e s were foun d.
In order to induce short-eulm mutation in
ri ce
V a r i e t y Norin No®22, i t s seeds were treated w ith r a d i o ­
active
on ly
P^2 by Kauai et ,|1,
(1961).
those p l a n t s were sel ect ed
be not notably d e c r e a s e d .
mutant types i t
In mutant g e n er at io n s
in which y i e l d
seeaied to
After o b t a i n i n g some r e q u ir e d
was f e l t that by r a d i a t i o n ,
a long culm
20
variety
its
can be changed to a short culm form without imp ai rin g
pr od uc ti vi t y#
Gopal-Ayengar jet a l .
(1969)
m o d i f i c a t i o n of the e f f i c i e n c y
(dES)
on pre-soaked r ic e
investigated
of d ie th y l
seeds.
the
s ulp hat e
Highest mutation rat es
were noted a f t e r 2 hour treatment of dry ri ce
seeds with
T . 0 * dES and s u b j e c t i n g 48 hours pre-soaked r i c e
to 0 , 2 $ dES for 2 hours*
sensitized
of
and
*t was observed that pra-soaking
the seeds to chemical
husk from the
seeds enhanced
treatments and removal
t h e i r r at e o f
germination
the e f f i c i e n c y of the chemical two-fold.
For i n d u c i n g
(1969)
useful mut ations i n r i c e
s o me
i r r a d i a t e d r i c e p la n t s at the growing
addition
visible
in
ch a ra ct er s and chemical
components,
useful
b las t resistance
a-nd high p r o t ei n co n t e n t ,
ch a ra ct er s l i k e
Sharma e_t a l , . ( l 9 7 4 )
v a r i e t y 3agannath
variation
Mutants were
ea rly m a t u r i t y ,
s ub je cte d the seeds of r i c e
and Rexero-Rll to EMS t re a tm en ts .
t r e a t e d po pu lat ion of v a r ,
In
3ag annath a high range of
for days to heading was noted and maximum
variability
was ob ta in ed
in 23 hr treatment.
g e n e r a t i o n s 11 e a r l y maturing mutants (5
and 6 from Rexero-Rll)
the mutants also
In
and Mg
from Jagan na th
were ob ta in ed which showed high er
than the pare nt v a r i e t i e s ,
quality.
stage in
inducing u s e f u l mutations in both
fo un d po s s e s s in g
yield
Tanaka
to i r r a d i a t i n g the dormant seeds1* I r r a d i a t i o n
proved e f f e c t i v e
of
seeds
from v a r ,
3a g an na th one
showed an improvement in
fine
grain
21
Tstikada ejb
physical
(1976)
s tu d ie d mu t ag en ic it y of
and chemical mutagens en r ic e p l a n t s *
aoras
Hi gh es t
f r e q u e n c i e s of ch lo rop hyl l mut ations and mut ations of
agronomic t r a i t s were obtained a f t e r
these
frequencies
from gamma r a y s ,
EMS treatments and
seemed to be three times more than those
Re garding mutagenie e f f e c t s ,
et hyl methane-
£0
su lp ho n ate and nitromethyl urea seemed e q u i v a l e n t to
gamma rays* D i f f e r e n c e s in mutation
between
spectrum were observed
gamma rays and chemicals and among c h e m ic a l s .
In order to study the freque ncy and spectrum of
ch l o r o p h y l l mutat ion s in r ic e i n
r e l a t i o n to the
and nature of the mutagen Bhan and Kaul
the
(1976)
genotype
s u b j ec te d
seeds of th ree r i c e v a r i e t i e s to gamma rays and two
a l k y l a t i n g agents
(EFIS and dES) alone and in
c om bi na tio ns .
O
They noted an enhanced chlorophyll mutation fr equency with
i n c r e a s i n g dose but the doses showing 90% s e e d l i n g
lethality
showed a drop in mutation frequency*
type ohlorophyll mutants c o n s t i t u t e d
a major c l a s s in
c h l o r o p h y l l mutants in M^ in both p h y s i c a l
treatments.
ENS was r es po ns ib le
Al bi na
and chemical
for in d uc in g
significantly
h i g h e r proportion of a l b i n o s than did gamma r a y s 0
N a i r and Ninan
fertility
u s i n g Co
(1977)
seed
and ch lo rop hyl l mutation frequency in rice
fin
gamma rays and EMS# A high er percentage of
chlorophyll
d e f i c i e n t mutants a f t e r EMS treatment and
a h i g h e r percen tag e of
observed.
s tu d i e d induced
s t e r i l i t y with gamma rays were
A dose dependent l i n e a r
i nc r ea se i n
higher
sterility
c l a s s e s was obtained
in both the mutagens*
In both mutagens the s t e r i l i t y
c l a s s e s below 5 0 % gave
the maximum frequency of ch lo rop hyl l
this
e f f e c t al so
conducted s t u d i e s on the
chromosome and chlorophyll
irradiation,. D ifferen t
loc us i n rice
response
to gamma
doses of gamma rays r e s u l t e d in
chromosomal a b e r r a t i o n s and with
radiation
d e f i c i e n t mutants;
proved to be dose dependent,
flajuradar ( 1 9 7 9 )
of
22
the i n c r e a s e of
dose an i n c r e a s e in number of ch lo rop hyl l
mutants was obs erv ed *
The t o ta l
number of ch lo ro ph yl l
mutants induced was hig he r as compared to the induced
fre que ncy of chromosomal a b n o r m a li t ie s *
In order to examine any d i f f e r e n c e s in the radioa e n s i t i v i t y of four in b re d l i n e s
701)
and two h y b r id s of maize
xN6)Notani
and
(1961)
(04,
(
D65,
N§ and
Ite
D 65 x 0 4 and I t e 701
irradiated their
dry seeds w it h X-rays
thermal neutron doses* D i f f e r e n c e s in r a d i o s e n s i t i v i t y
among inbred l i n e s as well as among h y br i ds were noted*
From the r e s u l t s i t
appeared that r a d i o s e n s i t i v i t y was
c o r r e l a t e d with growth rate of a l i n e evidenced
s e e d l i n g he ig ht*
from
F^ hy br id s were in general more r a d i o ­
r e s i s t a n t than the pa re n ts e s p e c i a l l y at high er
whic h was a t t r i b u t e d to the gene ral
doses
homeostatic and
h e t e r o t i c p r o p e r t i e s o f hy br ids *
Gardner ( 1 9 6 9 )
pollinated
initially
r esponse
sample
v a r.
Hays
s ub je cte d
seeds of corn
Golden) to thermal neutron
and a ga in at gen er ati on 2 .
to s e l e c t i o n was n eg a ti v e
u n t il
(open
in
treatment
It was noted
that
the i r r a d i a t e d
treatment was d i s c o n t i n u e d and from
23
g e n e r a t i o n 3 onwards a rapid l i n e a r response o f 4%
per
generation was ob s er ve d .
increased prolificacy
Thermal neutron
treatment
and thus in c r e a s e d a d d i t i v e
v a r i a n c e for both p r o l i f i c a c y
and grain yield,,
g en et i c
Selection
pressure
in both control and i r r a d i a t e d po p u la t io n s
resulted
in higher grain y i e l d ,
greater p r o l i f i c a c y ,
taller
p l a n t s and l a t e r m a t u r i t y ; however,
s e l e c t e d control
in
and i r r a d i a t e d
at gen er ati on 1 0 ,
s
populationA were equal but
case of i r r a d i a t e d po pulation
higher
se l e c ti o n
Si n g l e to n
good fu tu re g a i n s and a
< l i m i t were predicted',
(1969)
tr ie d to induce mutations in maize
w i t h thermal neutr ons u t i l i z i n g
the proposal
that mutation
can be ob ta in e d a f t e r one s e l f - p o l l i n a t i o n in maize i f
same mutated c e l l
ear Bind ca rrying
goes to make up both the t a s s e l
the m aterial
to the
the
and the
generation.
The r e s u l t s o b t a i n e d showed that 4 08 $ of the t o ta l
p r o g e n i e s tested
mostly
were
in
segregated for
good monogenic r a t i o ,
for albino
seedling,
c h a r a c t e r s as l u t e u s ,
3ordanka
(1978)
some
s e e d l i n g c h a ra c t e rs
Further most s e g r e g a t i o n s
but a few were for
ye llo u- gre en,
v ir e s c e n t
such
and d w ar f ,
studied the e f f e c t s of r e pe ate d EMS
and NMU treatments on the m u t a b i l i t y in C-103 and F-7 inbred
l i n e s of m a i z e .
mutant plant
Following repeated NMU treatment,
the
frequency and spectrum did not i n c r e a s e
compared to Q I S ,
Uith repeated EMS treatment o f
h e r e d i t a r y v a r i a t i o n was in d u c e d .
s ee d s ,
Ho we ver,repeated treatments
w i t h EMS and NNU proved of no s i g n i f i c a n c e
b r e e d i n g of m a i z e .
as
in mutation
24
P u l 88 8
Kl ing m ull er
radiation
(1961)
noted a marked i n c r e a s e in
s e n s i t i v i t y in V lc ia
fa ba minor when the
see ds were dr ie d to low moisture content bef or e
irradiation,,
W e l l e n s ie k
cultivar
pea
(1961)
irradiated
dry seeds o f pea
d o m i n a n t * « In the progeny of one i r r a d i a t e d
seed two m ut an ts,
namely * e a r l y
*v e r y ea rly f l o w e r i n g *
thB o r i g i n a l
f l owe rin g*
and
were found which d i f f e r e d
c u l t i v a r by 13 and 28
from
days r e s p e c t i v e l y
in
time of fl o w e ri n g *’
Gottschalk
(1961)
obtain ed ne a r ly 400 mutants in
P;isum through X - ir r a d ia ti o n o f i t s
Ahund-Zade
pe a var*
seedsV
and Hvostova ( 1 9 6 6 )
'Pioneer*
tre ated dry
seeds of
with vario us mutagens for c y t o g e n e t i c a l
a n a l y s i s of the mutagenic e f f e c t s of i o n i z i n g
and a l k y l a t i n g compounds.
Symmetrical
radiations
5
translocation,
i n v e r s i o n s and m ic r o n u c ls i were observed a f t e r m e io t ic
studies**
Reduced
seed set r e s u l t e d a f t e r treatment with
gamma rays and f a s t neutrons*
Variability
been
the
in
( V i c i a faba L * )
quantitative
cha rac te rs of
was s tu d i e d by D i s l e r
(1966)
i n f l u e n c e o f mutagenic f a c t o r s 1. An i nc r e a se
r an ge o f
spontaneous v a r i a b i l i t y
in plant h ei g ht
number of pods and seeds per pl a nt
was observed a f t e r
field
under
in the
and
in mutating c u l t i v a r
treatments wi th E l ,
D1S and gamma r a y s .
25
Moh ( 1 9 7 2 )
of
induced v a r i a t i o n s in seed coat colour
some black bean
( Phaseolus v u l o a r i s ) v a r i e t i e s of La t in
America* M u t a g e n i s i s r es ul t ed in
in d uc in g some seed coat
c o l o u r mutants which v a r ie d from w h i t e ,
yellow to vario us
d e g r e e s o f brown and t h e i r seed coat colour was a ss o c i a te d
w i t h a change in
hypocotyl colour from red to green*
t h e s e mutants were b ea r in g white f lo w e rs in ste ad
the pa ren ts but t h e i r morphology,
resistance
were s i m i l a r
N a r si ng ha ni
eou pea
In M^
of red of
growth h a b it and d i s e a s e
to those of the p a r e n t s .
and Kumar ( 1 9 7 6 )
( Viana s i n e n s i s L .
t re a tm en ts*
All
su bj e c te d the
Sav iex Hassk )
t© EMS and MMS
and Mg g e n e r a t io n s reduc ti on i n
p e r c e n t a g e , mean pod number,
seeds o f
su rv iv a l
seed y i e l d per p l a n t and
a v e r a g e po lle n f e r t i l i t y was observed which was l e s s in
Mg*
A few long podded mutants,
ch lo rop hyl l mutants and
l e a f l e t m o d i f i c a t i o n s were recorded*
Mujeeb and Greig ( 1 9 7 6 )
induced n a t a t i o n s i n French
be an
( Ph as eol us v u l o a r i s ) var*
with
gamma rays*
seed
g er m in a t io n ,
leaves*
However,
Irradiation
“ Blue Lake** by i r r a d i a t i o n
acted as a stimulant for
seedling height,
chloro phy ll
and leng th of primary
con tent per l e a f
area remained
unaffected*.
Prasad
(1976)
attempted v a r i e t a l
grain ( Viona r a d l a t a )
potential
and at the
improvement o f green
to i s o l a t e mutants having h ig h y i e l d
same time po s s e s s in g t o le r a n c e to
d ro ugh t c o n d i t i o n s p r e v a i l i n g i n
dry lands'* Pre-soaked
26
se e ds o f §reen gram var*
RS 4 were subj ec ted
to
aqueous s o l u t i o n s o f 0 , 2 $ and 0 , 3 $ c on c e n t r a ti o n s of
EMSy Ten mutants with higher number of pods than parent
were
scored in
ar,c* tested under dry land c o n d i t i o n s
a l o n g u i t h the pa ren t and the h i g h e s t y i e l d i n g
ch e c k ,
A mutant
s t r a i n M 8 was found c o n s i s t e n t l y m a in ta in in g i t s
superiority
to
in y i e l d and pod number per pl a nt as compared
the parent and h i g he st y i e l d i n g
c o n d i t i o n s of r a i n f a l l
of
and also
higher yield p o te n t ia l,
tolerance
exhibited
a combination
e a r l i n e s s o f maturity and
to drought conditions',
Nerkar ( 1 9 7 6 )
Lathvrus
check under vary ing
tr ea te d seeds from 5 v a r i e t i e s of
s at ivu s u i t h gamma r a y s ,
tre a tm en ts of gamma rays and NMU,
f r e q u e n c i e s of ch lo ro ph yl l
ENS,
NMU and combined
Af ter computing
and v i a b l e mutations on
progeny b a s is i t uas found that they were dose-dependent,
i r r e s p e c t i v e of the
genotype,
but while
calculating
f r e q u e n c i e s o f ch lo ro p h y ll and v i a b l e mutations on the
b a s i s of
all
^2
po p u la t io n t h i s trend was not ma in ta in ed
genotypes,' Nitrosomethyl
in
urea uas found to be the
most potent mutagen for the i n d u c t i o n of mutat ion s followed
by EMS and gamma rays*. The combined treatments re ve a le d
no
s y n e r g i s t i c effects*,
Rao and 3ana ( 1 9 7 S )
gram ( P h a s e d u s munoo )
w i t h the o b j e c t i v e
s ub je cte d the seeds of black
to X-rays and EMS treatments
of ob ta in in g
*
some promising m ut an ts .
27
The induced l e a f mutants scored comprised of c r i n k l e d
leaf,
waxy-leaf,
narrow-leaf and u n i f o l i a t e mutants*
V a s i l e v a and Mekhandzhiev ( 1 9 7 7 )
of
two pea c u l t i v a r s with 0*1
©f dES and EMS and 0 * 0 1
treated
- 0 * 6 $ buffered
dry seeds
solutions
- 0 . S # s o l u t i o n s of El * Mutagenic
a f f e c t was found dependent upon mutagen con ce nt ra ti o n and
plant
genotype*,
EMS treatments gave hig he st mutation f r e q u ­
ency w hi l e dES treatments the l o w e s t .
In
Phaseolus aureus Roxb*
r a y s on dry s e e d s ,
and
the e f f e c t of gamma
seeds pre-soaked i n d i s t i l l e d
water
seeds pre-treated wi th 0 , 2 $ c o l c h i c i n e was s tu di ed
by
Raghuvanshi er^ a l «, ( l 9 7 8 ) v Pre-soakirigof seeds was found
to
a f f e c t the germination a d v e r s e l y .
fringed
male
leaf,
strap l e a f ,
advanced
In M2 ge ne r at io n
stigma and t ri s o m ic and
s t e r i l e mutants were i s o l a t e d *
To t es t the e f f e c t i v e n e s s of gamma r a d i a t i o n on
the
i n d u c t io n of
varieties
q u a n t i t a t i v e v a r i a t i o n s i n 3 broad bean
( V i c i a fa ba L „ var*
Kasim .et a l J ( 1 9 7 8 )
ray
doses*
In M^
Sinjar,
exposed the
Egyptian and French)
seeds to d i f f e r e n t
gen er ati on response in
v a r i e t i e s was d i f f e r e n t but a ll
the three
the V a r i e t i e s e x h i b i t e d
e a r l i n e s s of f l o w e r i n g and i n c r e a s e i n stem l e n g t h *
all
gamma
In
the v a r i e t i e s te st ed no r e l a t i o n s h i p e x i s t e d between
th e magnitude o f i r r a d i a t i o n and the amount o f induced
genetic v a r i a b il i t y .
28
Molkhova and V a s il e v a ( 1 9 7 7 )
isolated
three
gamma ray indu ced mutants of pea#
Chandra and Tewari ( 1 9 7 8 )
bean
observed that in
( Phasealtis aureus Roxb) v a r .
S-® and Pusa B a i s a k h i
i n c r e a s i n g doses of gamma rays and neutrons caused a
gra dua l
ovule
leaf
re duction in germination of seeds and pollen
fertility.
I r r a d i a t i o n caused the appearance of
a b n o r m al it ie s i n c l u d i n g u n i f o l i a t e ,
trifoliate,
tetrafoliate
Bhat tac ha rya
va r*
bifoliate,
and p e n t a f o l i a t e
(1978)
noted that in
characters*.
Glycine max L .
Bragg a dose o f 10 Kr gamma rays improved
yield
and
and other ch a r a c t e r s while
growth,
higher doses showed
progressive in h ib it io n .
N i k n e j a d et. a l « ( 1 9 7 8 )
spectrum of a high ly
line
tic
t r i e d to a l t e r the mutation
spontaneous mutable chi ck pe a pure
( Clear arietintim L . )
by treatment with two radicmime-
substan ces - EMS and NaN^ but t h e i r attempt f a i l e d in
pr o d u c in g a s i g n i f i c a n t change in mutation spectrum.
In order to demonstrate the role of DMSO i n I n f l u e n c i n g
the mutagenic e f f e c t of EMS Chaturvedi
and Singh
(1978)
s u b j e c t e d dry seeds o f mung been cv# Pusa B a i s a k h i
aqueous
s o lu t io n s of mutagen and \ % DMSO.
indicated
to
S tu d i e s
that the DMSO enhanced the e f f e c t i v e n e s s of
ch em ic al mutagen which was e v id e n t from reduced
germination p e r c e n t a g e ,
and
seed f e r t i l i t y
se ed li n g
height, pollen
in M«j g e n e r a t i o n ,
fertility
and i n c r e a s e d
29
frequency,
P^2
spectrum of chloro phy ll
and vi a bl e mut ati ons in
generation.
Das and Prasad ( 1 9 7 8 )
differential
(1 0 - 5 0
Kr)
s t u d i e d the i n f l u e n c e of
and combined treatment of gamma rays
and 0 O2 $ methyl
methanesulphonate (PIES) on
some v a r i e t i e s o f La th vr us s a t i v u s L .
h e i g h t of the p l a n t and
showed doss dependent
It uas noted
that
number of branches per p l a n t
reduction
in a l l
the v a r i e t i e s
at M^ and both i n c r e a s e and decrease at Mg as compared to
control*
Khan ( 1 9 7 9 )
t r ea te d the dry seeds of mung bean
( P h a s e o l u s aureus R o x b . )
Mu tag en ic
var*
PS-16 u i t h va ri o us mutagens.
treatments r es ul t ed in a p o s i t i v e
s h i f t of mean
v a l u e s for a l l po ly g e n ic t r a i t s exc ept for p l a n t h e ig ht
and days to flower** Treatments also induced v a r i a t i o n s for
many q u a n t i t a t i v e
ch a ra ct er s which appeared important for
the improvement o f mung bean*
I r Mung bean
Hashim
(1979)
( Phaseolus aureus Roxb.)
s tu d ie d
the r e l a t i v e mutagenic e f f e c t i v e -
n e s s of three mutagens gamma r a y s ,
and h y dr az in e hydrate
Khan and
(HZ).
A
methanesulphonate
HZ proved an e f f e c t i v e
(EMS)
agent
fo ll o w ed by EMS and gamma rays when e f f e c t i v e n e s s was
measured as the frequency of mut ations induced by u n i t dose
o f mutagens*
30
Prasad and Prasad ( 1 9 7 9 )
combined e f f e c t s of
on
quantitative
stud ied
differential
a lk a ne su lph on ate s and a n t i b i o t i c s
ch a r a c t e r s of French bean ( P h a s e o l a s
v u l g a r i s )o
The p a t t e r n of e f f e c t produced by raono-
functional
a l k y l a t i n g agents in r e l a t i o n
variance
and
to mean, r a n g e
and co-variance was s i m i l a r to that of
Dry seeds o f
antibiotics^
red gram (C a i a n u s caian varV 3 5 )
were
s u b j e c t e d to d i f f e r e n t doses of gamma rays and d i f f e r e n t
c o n c e n t r a t i o n s of radiorairaetic substan ces l i k e
N-nitroso-N-methyl
EMS and
urea by Chaturvedi and Sharma ( 1 9 7 9 ) ,
A number o f mutants wi th good agronomic q u a l i t i e s was
scored and some of them which bred true i n
multiplied
and M^ were
farther*,
Raghuvanshi
and S i n g h
(1979)
demonstrated
the
d i f f e r e n c e s in r a d i o s e n s i t i v i t y
of
l o i d s in
T r i o o n e l l a feenum-oraecum L
at
different
40 Kr dose of
genotypes of
gamma rays*
autotetraploids of
d i p l o i d s and autotetrap-*
R es ul ts
showed th at even
d i f f e r e n t genotypes be lo ng in g
to same
s p e c i e s may show tremendous d i f f e r e n c e s in t h e i r r a d i s —
sentivity^4
To enlarge
early
the na t u r a l l y oc c ur r in g v a r i a t i o n s in
v a r i e t i e s of green gram ( Viona .radlata L . )
its
s e e d s were i r r a d i a t e d with gamma rays by Singh e_t _§1*
(1979)*
I t was seen that a d i f f e r e n t i a l
irradiation
doses e x i s t e d
no ted that a l b i n a ,
T
Ace Ko . X
response
among genotypes*
I t was fur th e r
xantha and c h l o r i n a types of
T
. I & 72. ......
r
Ik*.
,
to
ch lorophyll
31
m utations
segregated
in
tha
ratio
of
1 mutant
S 15
norm als*
Katyayani
at ,§,1.(1980 a)
e f f e c t s of maleic hy dr az id e
studied
(MH)
the mutagenic
and ethyl methane­
sul pho na te
(ENS) on germinating seeds o f T r i o o n e l l a foenum-
□raecum L .
Results
EMS (o.Q5- 0 . 1 % )
on
showed that high er c o n c e n t r a t io n s of
and MH ( 0 . 1 $ )
exercised
retarding effect
s e e d l i n g growth w h il e low co n c e n t r a ti o n s of both the
chemicals,
resulted
p a r t i c u l a r l y 0 . 0 0 1 $ EMS and upto 0 . 0 5 $
in i t s promotion.' It was also
tre atment with 0 . 0 0 1
i n d u c t i o n of e a rl y
and 0 . 0 1 $
MH
observed that seed
and EMS r e s u l t e d in
flowering^*
In Phaseolus aureus Roxb. var. Pusa Baisakhi radiation
induced mutagenic studies were done by Katyayani e_t al,.
(1980
b) after exposing its seeds to gamma rays'. Studies
revealed that germination percentage showed an increase
with lower doses of gamma rays ( 1 —7 Kr
), the
maximum
being at 4 Kr treated samples but with higher doses
(8-10 Kr) a gradual decline in germination percentage was
observed. The application of doses ranging from 5-7 Kr
caused an increase in plant height whereas doses from 8—10 Kr
induced early flowering and also increased number of pods
per plant.
32
Vegetables
In order to determine the r e l a t i v e mutagenic a f f e c t s
of
EMS and X-rays on ind uc tio n of mutations in
annuum L ,
its
Zubrzycki
seeds with EMS
and WoFr*de^ Pahlen ( 1 9 7 2 )
and X-rays*
Capsicum
t rea ted
A ft er a n a ly s in g Mg
p r o g e n i e s i t was noted that the chemical agent proved to
be wore e f f i c i e n t
and e f f e c t i v e
in the in d u c ti o n e f
c h l o r o p h y l l n u t a t i o n s whi le in r es p e c t o f i n d u c t i o n of
m or pho log ica l m ut ati ons both the agents proved e qu a ll y
effective*
I t uas also noted that the chemical
agent
cau sed more damage to evule v i a b i l i t y
than X-rays,
b a s i s o f number o f chimeras o b t a i n e d ,
it
that
g iv e
On the
uas concluded
at the time of treatment d i f f e r e n t meristems, which
rise
to f r u i t s ,
existed
To increase v a r i a b i l i t y
( Capsicum annuum )
s e e d s to several
Skr ipn iko va
in the
in
seed*
sweet peppier
(1976)
chemical mutagens*
s ub je cte d i t s
In M^ g en er at io n mutagenic
treatments r e s u l t e d in decrease of germination
survival
and with the inc re a se
and
in mutagenic treatments
r e t a r d a t i o n of p l a n t growth and development i n c r e a s e d
progressively,
Khuspe and Ugale ( 1 9 7 ? )
demonstrated the e f f e c t of
gamma rays and EMS on growth and f r u i t
Capsicum
annuum.
-------------
In both M1 and M_
z
development of
g en er ati on s f lo w e r in g
got delayed to a remarkable ex t e n t a f t e r mutagenic
trea tme nt s but EMS proved more e f f e c t i v e
in t h i s r e s p e c t .
33
The weight of 100
dry fruits increased following
radiation doses but EMS was ineffective in thi3 respect
b o t h in
and M^*
R a d io cy to ge ne t ic a l
ware done by K a t i y a r
s tu d ie s
(1978a)
in Capsicum annuum
after
subjecting i t s
dormant dry seeds to d i f f e r e n t doses of gamma r ays *
C y t o g e n e t ic a l
ing
s t u d i e s revealed raeiatie a b n o r m a l i t i e s i n c l u d ­
stickiness,
bridges,
clump ing ,
unequal
m ic r os po r es *
fertility
segregation,
associations,
breakage,
la g g a r d s and abnormal
Induced chromosomal ab no r m al it ie s
and p o l l e n
were found to be dose d e pe nd en t,
Katiyar (1 9 78 b )
reported an induced de sy na p ti a
b e h a v io u r in a v a r i a n t
of
alt er ed
is o l a t e d
from i r r a d i a t e d po pu lat i on
C . annuum f o l l o w i n g 20 Kr gamma i r r a d i a t i o n
phenomenon r es u lt e d
and t h i s
in the prod uct io n o f m e io t ic anomalies
such as rnicronuolsi,
polyspory and
gametes with varying
d e g r e e s of gametic imba la nce ,
Mal tse va ( 1 9 7 8 )
gamma i r r a d i a t i o n
stimulatory a ct io n of
on seeds of th ree pepper s p e c i e s using
a dose o f 120 G-I 800 r*
in
detected
The stimulatory e f f e c t was d is p la y ed
a more rapid development of p l a n t s ,
early ripening*
The e f f e c t s observed
earlier
a ft e r
f l o w e r i n g and
gamma i r r a d i a t i o n
were l o s t a f t e r about 5 or 6 ha rve sts *
In
(creole
Colombia,
potato)
Solanum p h u r e i a 3 u z ,
presen ted
at B«k
3 ome problems to farmers because
of la ck of dormancy and also to g e n e t i c i s t s and b re ed ers
34
because of gametophytic self-incompatibility,, In order
to overcome these problems Gomez Cuervo and Estrade
Ramos ( 1 9 7 2 )
tried to alter both the characters by
artificial induction of mutations* After seed as well as
tuber irradiation phenotypical changes observed in subsequent
generations proved mostly harmful for the plants.
However, in the irradiated progeny one mutant was isolated
whose tubers did not sprout for seven months and another
gave fairly high seed set after pollination within the
clone.
Bob and Demoura ( 1 9 7 6 ) s t u d i e d the e f f e c t s of low
dosage of gamma i r r a d i a t i o n
and
sugar and p r o t e i n
War*
Russel Burbank
potatoes
that
content of p o t a t o ,
and Nampa,
growth,
yield
Solanum tuberosum L .
A f te r exposure of
seed
to d i f f e r e n t doses of gamma rays i t was noted
higher
Decre ase d
on emergence,
doses of i r r a d i a t i o n
tuber y i e l d
s ug ar content of
delayed p l a nt emergence,
and tended to i nc r ea se r ed uc in g
tubers.
Total
sugar content was v a r i a b l e
de pe nd ing upon c u l t i v a r and cu lt ur e
used'.
An i n v e s t i g a t i o n was undertaken by Majid
(1975)
to study and compare the frequency and spectrum of
m u t a t i o n s induced by non-io nizin g r a d i a t i o n
an i o n i z i n g r a d i a t i o n
age nt
(EMS)
and an a l k y l a t i n g
in two v a r i e t i e s of Ly c o p e r s ic o n esculentum and
one v a r i e t y of L ,
physical
(Co^-gamma ray s)
(UV l i g h t ) ,
peruvianum.
Combined treatments of
and chemical mutagens were also te sted
fa r
a s c e r t a i n i n g the type of i n t e r a c t i o n caused by such
35
com bination*
Treatment u it h e i t h e r ph y s ic a l or chemical
mutagen uas not as e f f i c i e n t in producing a wide spectrum
a f mut ations as the combined treatments which proved
quite efficacious
in producing high er mutation frequency
w i t h broad spectrum of vi a b le m u t a t i o n s .
v i a b l e mutations a f f e c t i n g growth,
leaf,
flower and f r u i t s
habit,
I good number of
c h a ra c t e rs of
were s c o r e d , ( See also M a j i d , 1 9 6 9 ) ,
In a mutation b re ed ing programme undertaken by
Maltseva
( 1977a )
i t was noted that the action of pre-
i r r a d i a t i o n of tomato seeds with a stimulatory
of
gamma rays on the
growth,
fr u itfu ln e s s of plants,
It
dose (2 K r)
development fl o w e r i n g and
depended on t h e i r moisture con tent.
was noted that seeds at moisture content o f 35-
■ x h i b i t ed no sti mul ato ry e f f e c t ,
levels
showed a s l i g h t
seeds at 11-12 $ moisture
stimulatory e f f e c t and seeds at
7 - 8 $ moisture conte nt e x h i b i t e d
effect
40$
the g re ate st stimulat ory
after pre-irradiation.
In a s im ila r experiment M al t se va
(1977b)
fur the r
demonstrated the dose rate dependence o f stimulatory
a c t i o n of gamma r a d i a t i o n on tomato s e e d s . O b s e r v a t i o n s
revealed
that r a d i o b i o l o g i c a l
dose of 2 Kr on tomato
e f f e c t s of the optimal
seeds depended upon the dose r a t e ,
A dose r a t e of 70-300 r/rain was recorded to induce maximal
s t im ul a to ry e f f e c t ;
dose rates l e s s
than 70 r/ m in were
n o ted to induce a weaker stimulatory a c t i o n ;
1000
r /m i n a n d
high er
a dose of
doses e x h i b i t e d no stimula to ry e f f e c t .
36
Zagoreheva and 3ordanov ( 1 9 7 7 )
obtained
soma
m ut an ts of tomato a f t e r gamma i r r a d i a t i o n of s eed s with
d o s e s of 20 and 30
Kr.
m utants ( a t r i p l o i d
In the course of
study two genomic
a n d a t e t r a p l o i d form) and a
chromosome a be r ra t io n of the t r a n s l o c a t i o n
separated.
To determine
the
e f f e c t s of pre-sowing gamma
i r r a d i a t i o n of seeds on growth,
tomatoes ( v a r .
Revermum F)
type were
development and y i e l d of
Ge r t s u s k ii
mj± j|l.
irradiated
the tomato seeds with gamma r a y s .
a t 5 0 —3000
r was noted r e sp o n s i b l e
for
and development and incr ea se d y i e l d .
by
14~$6% r e su lt ed by i r r a d i a t i o n
also
r e su lt ed
tomato l i n e
results
Irradiation
a c c e l e ra t e d
growth
An i n c r e a s e in y i e l d
at 50 and 250
r which
in a 22-54% gain in e a rl y f r u i t yield,,
G o r d a n o v et al*
the
(1977)
(1977)
XXIV - a ,
s tu d i e d induced mut age nes is in
using gamma rays as mutagen.
showed th at with 20 a n d
Their
30 Kr doses the w ide st
spectrum of mut ations was produced and a dose s l i g h t l y
above 70 Kr proved
lethal*
A considerable
g e n e ti c
div ergence
i n Mg and M^ g en e r a t io n s uas obs erved* Mutant forms with a
high
s o l i d con te n t,
d e h i c e on outs ide
Among i r r a d i a t e d
exserted
stig ma,
and p l a n t s withou t
short an th e rs which
shoots were re co r de d .
po pu lat ion s some forms with genomic
m u t a t i o n s were also no te d .
Yashvir
(1977)
induced p o l y g e n ic v a r i a t i o n in
l e n g t h of Abelmoschus es cu le nt us
fruit
by the a p p l i c a t i o n of gamma
37
rays,
EMS and dES .
(except
fruit
In M^ gen er ati on a l l
treatments
18 h EMS treatment )c a u s e d a decrease in mean
length,
In M2 ge ner ation t h i s parameter decreased
w i t h in c r e a s e in
increased
dose.
In g e n e r a l ,
v a r i a t i o n in f r u i t length
at lou doses and d ecr ea se d at higher doses*
Effects of X-irradiation on physiological and
morphological variability in Abelmoschus esculentus were
studied by Rao and Rao ( 1 9 7 8 )
after exposing its seeds
to gamma rays0 Irradiation dose3 from 1-3 Kr were noted
to increase germination uhile higher doses decreased
it* A delay in germination a n d reduction in growth rate
were noted at doses of 8 to 10 Kr*
Ra d i o c y to g e n e t i c a l
s t u d ie s on cuc ur bi t L u f f a
a c u t a n o u l a were undertaken by K a t i y a r
and Roy ( 1 9 7 7 ) .
It
was seen that with i n c r e a s e o f i r r a d i a t i o n dose a c o n s i s t e n t
increase
occurs.
in the frequency of every type of abnormality
Significant
types o f induced
post-irradiated
d i f f e r e n c e s were observed i n d i f f e r e n t
a bn o r m al it ie s between i r r a d i a t e d
stored
see ds,
Dhopte and More ( 1 9 7 8 )
brinjal,
Cluster
S o l a n u m melonoena L ,
(PPC)
and
and
irradiate d air dried
var.Long Uhite,
seeds of
Pusa Purple
Man jr ig ot a with gamma rays and R^
g e n e r a t i o n was an alysed to know the change i f
any in
q u a l i t y of f r u i t with respe ct to crude pr ot e in c o t e n t ,
vi t am i n C ontent and t i t r a b l e
acidity.
d os e s of 4 0 , 60 and 80 Kr were l e t h a l
The i r r a d i a t i o n
to the v a r . Long White
38
and PPC, while a dose of 80 Kr uas noted to be lethal for
¥ar« Manjrigota. Resultg indicated that gamma
irradiation
increased crude protein content of fruits by 0 * 8 7 $ in war,
Manjrigota at 20 and 40 Kr doses while other doses
decreased it in PPC by less than 1 $ . Irradiation treatments
also increased vitamin C content of fruits that ranged
from 8 . 1 3 to 5 6 . 9 1 $ in Long White, 7 . 0 9 to 8 . 6 2 $ in PPC
and 1 0 . 6 2 to 2 6 . 7 7 $ in Manjrigota. Gamma irradiation in
general also increased the titrable acidity of fruits*
Khan ( 1 9 7 9 a )
var.
treate d the seeds of
Purple Beauty with gamma r a y s ,
(E M S )
and d ie th y l
sulph ate ( dES ) .
some mutants in
et hyl methanesulphonate
A fte r EMS treatment
ge ner ation produced 76-92 f r u i t s per
p l a n t a g a i n s t average number of 22
In 1^2 ge ner ation
Solanam melonqena
f r u i t s in c o n t r o l .
from combined treatment u i t h gamma rays
and EMS some mutants showed a s i g n i f i c a n t i n c r e a s e in
number of f r u i t s
fruits,
and one mutant among them produced 134
which was noted to b@ more than s i x times o f
a v e ra g e number produced by control p l a nt s 5* F u r th e r,
some mutants were found b e ari ng f r u i t s
in c l u s t e r s while
some e x h i b i t e d d i s e a s e r e s i s t a n c e *
Seeds of Solanum melonqena tfar.
American Wonder were
s u b j e c t e d to the treatments u i t h gamma r ay s,
for
the ind uc tio n of mutations by Khan ( 1 9 7 9 b ) .
generation
in
EMS and dES
some mutants had two types of f r u i t s
shape and s i z e .
In
differing
Combined treatments with gamma rays
33
and
in
EMS produced
some mutants with
shape and s i z e
fruits modified
and combined treatments with gamma
r a y s and dES r e s u l t e d in the pr od uct io n of some mutants
with higher yields
and a mutant with a cre epi ng h a b i t *
I n d u c e d d is e a s e r e s i s t a n c e was observed in
o f Mg gene ration
and a mutant produced 38 f r u i t s
a v e r a g e o f 18 f r u i t s per plant in con trol,
Khan,
1980,1981,
Medicinal,
t h e i r e f f e c t on i t s
su bj ec te d
gamma rays to
in v e s t ig a t e
on the growth and m e tab ol ic
and frequency and spectrum of
Irradiation
with
different
doses resulted^
d u p l i c a t i o n s and t r a n s l o c a t i o n s both at m i t o s i s
a s / w e l l m e io s is *
increase
t o ta l
Uith i n c r e a s i n g
in seed l e t h a l i t y ,
Ohimdsal e f f e c t s
observed.
of
dormant seeds of
the production of chromosomal a b e r r a t io n s i n c l u d i n g
deletions,
of
s ee d s ,
of the p l a n t
v i a b l e mutations*
as
also
ornamental and other p l a n t s
D a t u r a metal to d i f f e r e n t doses of
in
(s e e
against
1 98 2a and 1 9 8 2 b ) *
Kaul and Si ng h ( 1 9 7 2 )
activities
some mutants
doses of r a d i a t i o n s
s e e d l i n g i n j u r y and production
such as morphological
f r e a k s were
In M^ p o p ul at io n an i n c r e a s e in the conc ent rat io n
a l k a l o i d s was noted which was more at lower
dose o f i r r a d i a t i o n ,
Kaul and Choudhary ( 1 9 7 2 )
Atropa b el lad on na
to d i f f e r e n t
exposed the seeds of
doses of gamma r a y s 0 S tu d i e s
were aimed at a s s e s s i n g the v a r i a b i l i t y
in po ly g en ic
40
c h a r a c t e r s releas ed
in
and Wg g en er ati ons *
v a r i a b i l i t y was noted in Pl2 g e ne rat io n than
observing
alkaloid
a greater v a r i a b i l i t y
content than that for
l e n g t h i t was i n f e r r e d
respond
for
differently
a wide
plant
that d i f f e r e n t
•
A ft er
number and
height and l e a f
ch a ra ct er s may
to d i f f e r e n t mutagenic treatments*
Mentha a r v e n s i s
to ob tai n
tiller
& hig he r
Kaul and Kak ( 1 9 7 2 )
were able
spectrum of mut ations p a r t i c u l a r l y with
ch emical mutagens though the number o f mutants was not h i g h ,
Dnyansagar and
of
diploid
Kothekar ( 1 9 7 9 ) ,
v a r i o u s morphological
diploid
the seeds
and t e t r a p l o i d p l a n t s o f Solanum nigrum L*
d i f f e r e n t doses of gamma rays to
studied,
su bjected
variable
characters.
to
study t he ir e f f e c t on
On the b a s i s o f parameters
response to gamma rays was noted between
and t e t r a p l o i d pl a nt s and the d i p l o i d Solanum njorum
proved more s e n s i t i v e
to gamma i r r a d i a t i o n
as compared to
the t e t r a p l o i d .
Broertjes
(1976)
irradiated
the c o l c h i c i n e
induced
a u t o t e t r a p l o i d s of
three Achimenes c u l t i v a r s w it h X-rays
and f a s t ne u t r o n s .
Re su lt s of the experiment in one of the
c u l t i v a r s were compared with the i r r a d i a t i o n
diploid
in
form and i t
results of
was noted that the mutation frequency
i r r a d i a t e d p o p u la t io n of the a u t o t e t r a p l o id was 20~40
times hi g h er than in the corresponding d i p l o i d *
wi th
a
some good agronomic
irradiated population.
Mutants
q u a l i t i e s were i s o l a t e d from
41
An attempt uas made by Abraham and De sai
to induce
variability
mu tan ts o f s i n g l e
and to o b ta in
(1976)
u s ef ul
some
and double type c u l t i v a r s of tuberose
( P o l v an th es tuberosa L * ) *
Thi rte en mutants were i s o l a t e d
among which eleven with l e a f colour V a r i a t i o n s belong ed to
single
type a n d two t o
double*1 The mutants appeared more
f r e q u e n t l y with f a s t neutrons than with X-rays and gamma
r ay s*
Tarar and Onyansagar
(1978,
comparative mutagenic e f f e c t s of
Turnera u l m i f o l l a L i n n ,
var.
198 0)
the
EMS and gamma rays in
anoustifolia U illd ,
r ec or de d varying de grees of meio ti c
t re atm en ts with the mutagens.
studied
They
irregularities
An in c r e a s e
after
in t h e i r f r e q u e n ­
c i e s was observed wit h the higher exposure to gamma rays
and hig he r co n ce nt ra t i o n and d ur ati on of EMS tre atments*
It was further noted that frequency of meiotic abnormalities
uas less in second generation whereas in third generation
they were noticed only occasionally* Plants raised from
irradiated or EMS treated seeds exhibited earliness or
delay in flowering, reduction in the size and number of
flowers, and variations in floral parts'* However, floral
abnormalities were more pronounced in the plants raised
from EMS treated seeds and appeared predominantly in the
M^ generation but rarely in subsequent generations'*
Prasad and Singh ( 1 9 7 8 )
saf
flower var. I . C .
irradiated dry seeds of
11842 with 45 Kr gamma rays from
42
a Co^
source,, One mutant gave a 6 3 , 8 $
y i e l d per pl a nt over
increased
the parent v a r i e t y ,
Behera a n d Pa t n a i k
exposed dormant seeds
(1979)
Amaranthus hypochondrlacus L ,
to d i f f e r e n t doses of
gamma rays and chemical mutagens (EMS and d E S ) ,
o f v i a b l e mutations a f f e c t i n g
u a s noted in
of
floral
and
an attempt to obtain
other
scor ed,
but
r e s p e c t of height
in
diploid
of
the crops
"G r o s M i c h e l " Va le z
its
rhizomes
deformations a n d
the
and small mor phological
in
and f r u i t c h a ra ct er s *
the v a r i a b i l i t y
for branc hin g
t o r ia to incr ea se the o i l
Gupta and Singh ( 1 9 7 4 )
s t r a i n of t o r i a
su bjected
( B r a s s i c a camoes tri s)
to gamma r a y s .
and treatment e l i t e s
populations
irradiated
they uere comparable to the control
and t e t r a p l o i d
autotetraploid
resistant
In M^ ge ner ation tuo mutants
pigmentation
In order to en la r ge
control
var*
one u i t h d r a s t i c l e a f
uith intensive
differences,
T-12
some d is ea se
and Cedeno Maldonado ( 1 9 7 2 )
u i t h 2 05 - 40 Kr gamma r a y 3 ,
ue r e
the le a v e s and i n f l o r e s c e n c e
in d i f f e r e n t pa tt er n s uas common.
mutants in Musa 3apientum L .
Fortune
A number
ge ne r at io ns among uhich f a s c i a t i o n
ax is r e s u l t i n g
In
seed
yield
s esd s of
and i t s
O b s e r v a t i o n s on y i e l d of
r eve al ed
gave high er y i e l d s
that all
the
treatment
than control1,*1 A s i g n i f i c a n t
d i f f e r e n c e uas noted betueen d i p l o i d and t e t r a p l o i d treatment
elites.
43
Rai
Linum
w it h
in
and Das ( 1 9 7 5 )
usitatissimum L .
irradiated
var*
Hira,
dry seeds of
Mukta and Neelam
gamma rays to induce mutations,.
different
types o f mutations in
U i t h an inc re ase i n
increase
Irradiation
re su lt e d
different varieties*
the dose of i r r a d i a t i o n an o v e r a l l
in mutation frequency was noted*
A considerable
i n c r e a s e was obs erved in a ll the mutants in t h e i r
flowering,days
days to
to m a t u r i t y 9 number of non-bearing t i l l e r s
p er p l a n t and number of le a ve s per cm on the main t i l l e r
while
a d r a s t i c re du ct io n was noted in t he ir p l a n t h e i g h t ,
number o f ca p su le s per pl a nt and number of
seeds per c a p s u l e .
In a gamma i r r a d i a t e d p o p ul at io n of peanuts
hvpoQaea )
M ou li
a suppressed branched mutant was i s o l a t e d by
and P a t i l
altered
( Arachis
( l 9 7 6 ) f*! The mutant had la rg er
fl ow e ri ng p a t t e r n ,
reduced
shelling,
leaves,
smaller kernel
and branch len g th b ei ng only 2 0 $ and 5 0 $ of normal
ir» the
autumn and spring growing seasons r e s p e c t i v e l y .
Seetharam and S r in i v a s a c h a r
s ee ds o f two
to
linseed varieties
gamma i r r a d i a t i o n of
irradiation
effects
(197S)
subj ect ed
( Linum usitatissimuni )
varying doses'*
In
hampered growth and s u r v i v a l ,
being pr o p o rt io n a l
the
to the dosage*
ge ne r at io n
the d e l e t e r i o u s
Irradiation
also
r e s u l t e d in the appearance of morphological a b n o r m a l i t i e s
and
delayed f lo w e r in g
in
In the
gen er ati on the 60 Kr
dose gave maximum frequency of ch lo rop hyl l
m ut a t io n s *
Among the chlo rophyll
and v i a b l e
mutations s c o r ed ,
44
viridis occurred more frequently. The visible mutants
scored included change in flower and seed colour, change
in petal arrangement and male and female sterile types.
In the selection of apple trees with good agronomic
traits Kolontaev and Kolentaev (197?) used radiation
mutagenisis.
Grafts of Anatonovka applta traes uere
X-irradiated and the three groups of morphological mutants
scored included slow, average and rapidly growing plants.
It was noted that after a dose of 6 Kr weak-growing mutants
appeared frequently while early growing ones appeared after
a 2 Kr dose. Each group of mutants differed in early and
late fruiting?type of fruiting and yield.
With the objective of obtaining some dwarf apple
mutatnts Kopan and Kopan (197?) subjected apple seeds to
gamma rays and chemical mutagens. Some treatments uere found
to be responsible for variability of hybrid seedlings in
relation to growth vigour and viability.
Chopra and Singh (l97s) studied the morphological
responses of Gujzotia abyssinica an oil seed crop of
economic importance, to three different concentrations
of 2, 4-D, 4 acute doses of gamma rays and varying combina­
tions of 2,4-0 concentrations and gamma ray doses giyen at
the seed stage1. Results indicated that combined treatments
exercised a greater deleterious effect on germination as
compared to individual treatments. Further it was noted
45
th at
grouth of the p l a n t s uas in v e r s e l y pr op o rt io na l
to the
but
increasing
gamma ray
doses and 2,4-D c o n c e n t r a t i o n s
g re a te r r e t a r d a t i o n uas recorded i n combined trea tments.
Treatments also induced
an omalies
a variet y of morphological
in leaves'*
Variability
in
the grouth o f
apple
tree
grafts after
g g
spring
treatment o f
sc io ns u it h
Co
gamma i r r a d i a t i o n was
s t u d i e d by Potapov and Kanashina ( 1 9 7 9 ) .
Acute i r r a d i a t i o n
r e s u l t e d in
reduced grouth
the production of forms ui th
characteristics.
Irradiation
also
r es u lt e d in
in c r e a se d bud
awakening and spur formation©' I r r a d i a t i o n of g r a f t s uith
f
1 . 5 to 4 . 5 Kr r e s u l t e d in grafts u i t h maximum spur f or m ati on .
2 .2 .
INDUCTION
OF
Das and Krishnasuami
se eds
(varieties
uas observed
in
t r e a te d
(1964)
su bj e c te d the mulberry
SRS -1 and Bush flalda )
c o n c e n t r a t i o n s of
It
POLYPLOIDY
colchicine
s o l u t io n
to d i f f e r e n t
to induce p o l y p l o i d y .
that percentage of germination uas higher
seeds than in control but higher co n c e n t r a ti o n s
proved l e t h a l to the
s e e d l in g s of both the v a r i e t i e s .
Immediate e f f e c t s of
treatment uere the a rr es te d grouth of
seedlings,
cotyledons and
thickened
su oll en h y p oc o ty l s.
U i t h the aim of producing t e t r a p l o i d s of mulberry
uith
higher y i e l d ,
for
3ilkuorm f e e d i n g and for
a source of breed ing ma ter ial
for
se rving as
the production of
triploid
46
varieties,
the
colchicine
for
t e t r a p l o i d s by
s e e d l i n g s with 0 . 4
and
d i f f e r e n t d u r a t i o n s . Maximum number
t e t r a p l o i d s uas recorded uith 0 . 4 % c o l c h i c i n e
with
a treatment time of 9 ho ur s .
buds u i t h 0 . 2 % c o l c h i c i n e
In
obtai ned
treatment of young mulberry
0.6%
of
Das e_t ai,0 ( 1 9 7 0 )
s olu ti on
s olu ti on
A 4 hour treatment of
also induced te tr a p l o i d y ,
ind uce d t e t r a p l o i d s gigantism uas observed in l e a f ,
floral
parts,
po ll en
and stomata;
s u c c u l e n t and more p u b es ce n t;
l e a v e s uere d a r k H j r e e n ,
po lle n
fertility
and
seed
s e t t i n g uere l o w e r .
S ing h and Kaul
polyploidy
ethyl
fo r 45 m i n u t e s .
possessed vigorous growth,
c o n t e n t and la rg er
seeds u i t h 5%
The po l yp l oi d s thus
increased a l k a l o i d
number of seeds per plant as compared
the c o n t r o l .
Ghosh ( 1 9 6 7 )
in
uere s u c c e s s f u l in i n d u c i n g
in Datura i n n o x ia by t r e a t i n g i t s
alcohol at 3 5 ° C
obtained,
to
(1967)
was s uc ce ssf ul
two varietiesjof r ic e
* Ashkata*
in i nd uci ng p o l y p l o i d y
and , D u l a r * ,
shoot immersion method under p a r t i a l
s e e d l i n g s possessed comparatively
thicker,
vacuum.
a tt ri b u te d
treatment r e s u l t e d in the
to the s tr uct ur al
a p i c a l meristem of r i c e „
in r i c e
and
Results ob ta i ne d
o f m ix o p l o id s in s t e a d of true c o l c h i p l o i d s ,
being
The tr ea te d
shorter height
stouter and darker-green l e a v e s .
shoued that c o l c h i c i n e
by employing
formation
the formation
c h a r a c t e r i s t i c s of
I n v e s t i g a t i o n s also revea led
that
there occurs diminution in vigour ui th p o l y p l o i d y .
Dnyansagar and Mhaske
of
Sida
of
colchicine
usual
like
for
seeds
12 and 18 hours r e s p e c t i v e l y ,
and
Induced t e t r a p l o i d s e x h i b i t e d
giga s c h a r a c t e r s a s s o c i a t e d with a u t o t e t r a p l o id y
larger
stomata,
sturdier plants,
flowers,
pollan
and l a r g e r
seeds and
Meioti© s t ud ie s d is p l a y e d u n i v a l e n t s ,
teivalents,
occasionally
diakinesis
and metaphase I ,
fibre
su bjected
rh om b ifo lia to 0 . 4 $ and 0 , 5 $ aqueous s o l u t i o n s
o b t a i n e d t e t r a p l o i d plants®
the
(1974)
o f the induced
t ha t of the d i p l o i d s
trivalentg
Fibre
and q u a d r iv a l e n t s at
s t u d i e s r ev ea led
that
t e t r a p l o i d s was stronger than
and i t s strength
reached the value of
t h a t of j u t e ,
Uith the o b j e c t i v e o f producing t e t r a p l o i d s of
some
s p e c i e s o f Zinnia which would prove of be tte r ornamental
value
Gupta and Koak
(1976)
t rea ted the seeds of
u i t h d i f f e r e n t co n c e n t r a t io n s of c o l c h i c i n e .
Z ael e g an s
Effective
c o n c e n t r a t io n s e x e r c i s e d
adverse e f f e c t on s u r v i v a l
plants.
Po ly p lo id s were,
therefore,
classes
having low s ur viv al v a l u e s ,
M o s a s h v il i
a v a i l a b l e only in
and Chankotadze ( 1 9 7 ? )
s e e d s with c o l c h i c i n e
under d i f f e r e n t
at
3 developmental
stages,
0,1$,
of
c o l c h i c i n e proved e f f e c t i v e
of the
tre ated
grape
c o n d i t i o n s of exposure
0 , 2 $ and 0 , 3 $ c on c e n t r a ti o n s
as i n d i c a t e d by the h i g h e s t
p e r c e n t a g e of p o l y p l o i d changes in root meristems,
Also
t hr ee
as a
t e t r a p l o i d s were i s o l a t e d
from the
seedlings
r e s u l t of c o l c h i c i n e treatment.
For the i n d u c t io n
of t e t r a p l o i d y
in C r o t a l a r i a brownei
48
and C ,
sericea
Gupta and Gupta ( 1 9 7 7 )
apices uith 0 . 2 $ colchicine
in
gene ral
solution.
uere found i n f e r i o r
treated
their
shoot
Induced t e t r a p l o i d s
to t h e i r
corresponding
diploids,
Uanjari
and Phadnis ( 1 9 7 7 ) s ub je cte d
Momordiea ch ara nt ia L .
colchicine
to d i f f e r e n t
to induce p o l y p l o i d y .
co n c e n t r a ti o n s of
Induced
d e v o i d o f vigorous grouth and e x h i b i t e d
Houever,
t h e i r fol
iar
and f l o r a l
the seeds of
t e t r a p l o i d s uere
delayed
flowering*
s i z e was b i g g e r
than
t h a t of diploids'*
I n d i r a and Abraham ( 1 9 7 7 )
reported r a d i a t i o n induced
t e t r a p l o i d in Capsicum annuum L 0 The t e t r a p l o i d
c h a r a c t e r i s e d by stunted grouth,
larger leaves uith larger
and number o f f l o r a l
c o m p l e te ly
sterile
f r u i t s be ing devoid
in 0 . 0 5 $
tion,
presence of t h i c k e r and
stomata and an i nc r ea se
organs.
in s i z e
In duced p o l yp l o i d uas
u i t h extremely poor f r u i t i n g ,
the
of seeds!,
Raut and Thorabre ( 1 9 7 7 )
tetraploidy
uas
were
s ucc ess ful
in
in d u c i n g
in Im pa t ie ns balsaraina L c by soaking i t s
and 0 . 1 $ c o l c h i c i n e ’.
In c o l c h i c i n e
p l a n t s shouing arr es ted gro ut h ,
seeds
t r e a t e d po pula­
h e ig ht
retardation^
abnormal plumules and l e a f shape uere o b s er ve d ,
A'utotetraploids po sse sse d big g er
blooming p e r i o d .
flouer
s iz e and longer
49
Semeniuk ( 1 9 7 8 )
induced p o l y p l o id y in the
f l o w e r i n g pot p l a n t Exactim a f f i n e
not h o r t i c u l t u r a l l y
Colchicine
were
Balt*
superior to o r i g i n a l
summer
T e t r a p l o i d s were
varieties.
ind uce d p o l y p l o i d s in flpluda mutica L .
reported by Flurty and S at yav ath i
(1978)*
Induced
t e t r a p l o i d s e x h i b i t e d robust s t a tu r e but the induced
d e c a p l o i d was reduced than i t s p a r e n t ,
Bakulin
Pooulus
(1978)
induced p o l y p l o i d y in d i f f e r e n t
species by s u b j e c t i n g t h e i r
c o l c h i c i n e treatments,.
s e e d l i n g s to
Induced p o l y p l o i d s e x h i b i t e d
d i s t i n c t i v e morphological p e c u l i a r i t i e s
st om ata !
some
and had la rg er
c e l l s and g re a te r number of e h l o r o p l a s t s than
diploids,
Roy and Mish ra
in
(1979)
while
in du ci ng t e t r a p l o i d y
Pha se olu s a c o n i t i f o l i u s noted that only the c o l c h i c i n e
treatment to a p ic a l
loids
parts.
buds proved
successful*
The autotetrap-
thU3 obtained e x h i b i t e d gigantism only in v e g e t a t i v e
C y to lo g ic a l
associations
ob se rv a tio ns r e v e a l e d va ri o u s chromosomal
such as u n i v a l e n t s ,
qu a d r i v a l e n t s o
bivalents,
In t e t r a p l o i d s normal
triva lents and
po ll en f e r t i l i t y
was f oll ow ed by normal f r u i t and seed s e t . By r a i s i n g
p l a n t s from these
seeds,
the
a re ve rs a l of t e t r a p l o i d s to
d i p l o i d s was o b s e r v e d .
In
an attempt to produce new forms of Nemesia strumosa
with l arg e
flo we rs and of a shorter h e i g h t ,
Gospodarek and
50
Huleuicz
its
(1979)
t r i e d to obtain p o l y p l o i d s by d o u b l i n g
chromosome 'mutnber using c o l c h i c i n e .
obtained
showed that though c o l c h i c i n e
r e s u l t in
chromosome doubling in
t h e i r morphological
The r e s u l t s
treatment did not
the c u l t i v a r s u s ed ,
cha rac te rs got changed,
namely,
the
p l a n t s uere shorter and t he ir f l o u e r s uere l a r g e r uhich
u e r e thought to be
the consequences of gene m u t a t i o n s ,
D y ! 3 e_t a l . ( 1 9 7 9 )
tre ated a p i c a l meristems of
P o r t u l a c a o r a n d i f l o r a ui th c o l c h i c i n e and uere
in
inducing polyploidy.
restricted
grouth,
m or p h o lo g ic al
studies,
Most of the treated p l a n t s
dark-green and t h i c k e r l e a v e s and
changes in the flowe r
those
differ
structure.
During
uhich had very thi ck l ea ves and uere
thought to be high p o l y p l o i d s .
flouers.
shoued
some p l a n t s a f t e r treatment produced no f l o u e r s ,
especially
gig a nt i s m
successful
I t was fur the r noted that
was much more pronounced
in l e a v e s than in
In induced p o ly p lo id s male f e r t i l i t y
from that of
l o w e r due to a very
diploids,
but
did
not
seed set uas three times
small number of ov ule s •