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 •
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