OOOO oO Backcross Method INTRODUCTION A cross between a hybrid (F; or a segregating generation) and one of its parents is known as backcross. In the backcross method, the hybrid and the progenies in the subsequent generations are repeatedly backcrossed t0 one of the parents of the Fy. As a result, the genotype of backcross progeny becomes increasingly similar to that of the parent to which the backcrosses are made. At the end of 6-8 backcrosses, the progeny would be almost identical with the parent used for backcrossing. The objective of the backcross method is to improve one or two specific defects of a high yielding variety, which is well adapted to the area and has other desirable characteristics. The characters lacking in this variety are transferred into it from a donor parent without changing the genotype of this variety, except for the genes being transferred. Thus the end result of a backcross programme is a well adapted variety with one or two improved characters. For example, a wheat variety, e.., Malviya 12, may become susceptible to a disease, e.g, leaf ris, but isan otherwise highly desirable variety. Ths variety, known asthe recipient parent, ig crossed tos leaf rust resistant variety, the donor parent, e.g. Sparrow. The Fy and the progeny in the catva ott generations are backcrossed tothe recipient parent, i.e, Malviya 12. During ome rust resistance is maintained by strict selection. In the end, rust recistan self-pollinated and homozygous rust resistant progenies are isolated. The same as the recipient parent, i.e., Malviya 12, except for their ru, susceptibility to rust of the recipient variety). Since the recipient py backcross programme, itis also known as the recurrent parent. The d known as the nonrecurrent parent because it is used only once producing the F,. 18.2. REQUIREMENTS OF A BACKCROSS PROGRAMME For the successful development of a new variety through the backcross requirements must be fulfilled. 1. A suitable recurrent parent must be available, which lacks in one oF two chara the backcross programme, t plants from the progeny are Se Progenies will be almost the St resistance (as compared to paren 4S repeatedly used in the on or Parent, ‘On the other hand, is ie breeding Programme for Method, the following a icteristics, 2. A suitable donor parent must be available. The donor parent should have the eh, ‘ter or the characters to be transferred in an intense form.Backcross Method 263 ie 3. The character to be transferred must have high heritability and, preferably, it should be determined by one or few genes, 4. A sufficient number of backcrosses should be made so that the genotype of the recurrent parent is recovered in full. Ordinarily, 6-7 backcrosses are sufficient for the purpose. In Practice, 4-6 backcrosses are usually made, 18.3. APPLICATIONS OF BACKCRoss METHOD SSS MNMSACK CROSS METHOD ‘The backcross method has been commonly used for the transfer of disease resistance from one variety into another. But it is also suitable for the transfer of other character, even quantitative characters, and is applied (o both self- and cross-pollinated crops. Backcross method may also be modified to allow the recovery of transgressive segregants, 18.3.1. Intervarietal Transfer of Simply Inherited Characters Characters governed by one or two major genes, e.g., disease resistance, seed colour, plant height, etc., are the most suited for transfer through the backcross method from one variety of the same species into another. Disease resistance has been the most commonly transferred character. Mostly such transfers are successful, but sometimes they may fail due to a tight linkage between the gene being transferred and some other undesirable gene; this phenomenon is known as linkage drag. 18.3.2. Intervarietal Transfer of Quantitative Characters Quantitative characters may also be transferred from one variety into another if they have high heritability. The following quantitative characters have been transferred using backcross method: earliness, plant height, seed size and seed shape. 18.3.3. Interspecific Transfer of Simply Inherited Characters Backcross method has been used to transfer simply inherited characters, mostly divease resistance, from related species into cultivated species. For example, transfers of resistance to wild fire-black fire from Nicotina longiftora to N. tabacum, of leaf and stem rust resistance from Triticum timopheevii, T. ‘monococcum, Aegilops speltoides and rye (S. cereale) to T. aestivum, of black-arm resistance from Several Gossypium species to G. hirsutum, etc, have been successful. Interspecific transfer of genes is ¢asy when the chromosomes of the two species pair regularly, But often chromosomes of the concerned Species are differentiated by structural changes that reduce pairing between them. As a result, in interspecific gene transfers often undesirable linked genes are also transferred along with the desirable gene, Another difficulty in interspecific gene transfers is that the transferred gene may not be able to function in the same way in the genetic environment of the new species, 18.3.4, Transfer of Cytoplasm Backcross method is used to transfer cytoplasm from one variet; cytoplasm is particularly desirable in cases of cytoplasmic or Variety or species from which the cytoplasm is to be transfe1 ackcross programme. The recurrent parent, the parent to which the cytoplasm is to be transferred, is teed as the male parent in the original cross as well as the backcrosses. After 6-8 backcrosses, the porgity would have the nuclear genotype of th recurrent parent and the cytoplasm fran gn, don Parent. For example, transfer of 7. timopheevii cytoplasm to T. aestivum hae resulted in cytoplasmie male sterile lines of T. aestivum. ‘ytoplasmic 'y oF species to another. The transfer of cytoplasmic-genetic male sterlity. The tred is used as the female parent in thePlant Breeding : Principles and Methogs a EES and Methogs .5. Transgressive Segregation os may be modified to obtain transgressive peepee tie meted ey be mov following two ways. First, the F, may be backcrossed only 1, St 2 times tp scared pan leaving much heterozygosity for transgressive segregants 10 Appear. In the seat tmodieation, tw ipa eaeninn parents may be used inthe backcross programme to acum Sa ta elit the backcross progeny. Such a modification ofthe backcross Would progus® oh vey that would not be exactly like any one of the recurrent parents. Hie iy 18.3.6. Production of Neamtsogenic Lines Near-isogenic lines are identical in their genotype, except for one gene. Such lines at us studying the effects of individual genes (and genes tightly linked with them) on yield ang ‘characteristics. Near-isogenic lines are easily produced using the backcross method (Chapter 7), ful Othe 18.3.7. Germplasm Conversion In some crops, valuable germplasms can not be utilized in breeding programmes since these lines ae Photosensitive and, as a result, do not flower under the prevailing photoperiods. Such lines may be used aoc urrent parents in separate backcross programmes with a photoinsensitive nonrecurrent arent in Alrica are disease resistant and possess excellent grain quality, butay Photosensitive, Consequently, they fail to flower in India S ‘ome of these land-races have been converted into photoinsensitive lines at ICRISAT, Hyderabad, these Converted lines can now be utilised in different breeding programmes, 18. GENETIC CONSEQUENCES. OF REPEATED BACKCROSSING 18.4.1. Reduction in Heterozygosity Repeated backcrossing leads to an incr a single gene A/a. Selfing of the F, heterozygotes and 1/2 homozygotes. and/2Aa, that is, 1/2 homozygotes at ease in homozygosity at the 1 (Aa) would produce 1/4 Backcrossing the F, ind 1/2 heterozygotes, In Same rate as does selfing. Consider AA, 12 Aa and 1/4 aa, i.e., Wt to the parent AA would produce 1/2 AA genes in a backcros: [2"-1/2"J" that is applicable to selfing. Thus, the first effe increase in homozygosity and in the frequency of homozygons 18.4.2. Increased Similarity with the Recurrent Parent In the backcross progeny (1/2 AA : 1/2 Aa), the frequency of homozygotes for the allele from the recurrent parent is 1/2, while in F, (1/4 AA : 1/2 Aa: 1/4 aa) its only 1/4. As the number of genes th! are segregating increases, the frequency of the desirable homozygotes, ie, of the recurrent parent YP decreases much more rapidly in the case of selfing c than it does in the case of backcrossing. i " number of genes, this frequency is (1/2)" in the case of backcross Drogen, but only (18) inthe progeny. Thus it is far more easier to sele irrent parent from a backe?® et for the genotype of the recui progeny than from a selfed progeny.' a a Backcross Method 265 ——“_<=-_-"*# its: When backcrossing is repeated, the frequency of desirable homozygote increases rapidly (Fig. 18.1) and in the sixth back-cross progeny, . ore than 98 per cent of the plants will have the AA genotype. The same is true for the other gene. -f the recurrent parent. Therefore, the genes from nonrecurrent parent are rapidly replaced by those 1:om the recurrent parent. The rate of this replacement is the same as the rate of increase in the Frequency of AA genotype. As a result, the genotype of backcross progeny becomes increasingly similar to that of the recurrent parent. Consequently, after 6 backcrosses about 99 per cent of the genes of backcross progeny would be from the recurrent parent (Table 18.1). This is expected when no selection is done in the backcross progenies. But often selection would be done for the plant type of the recurrent parent. This will result in a faster rate of return of the backcross Progenies to the genotype of the recurrent parent. HYBRIDIZATION NON-RECURRENT RECURRENT PARENT xX PARENT aa AA A Aa x AA BC, (Aa, AA x AA 50% ay (a, AA) x AA oo 25% 75% Ai (Aa, AA) x AA co 12.5% oe BC, (aa, AA) x AA 6.25% nee (a, A x “A 3.125% en (Aa, AA) x “A 1.562% ea BC, BC, BC, (la, AA x AA BC, 0.781% mA (a, AA) x AA 0.391%" ot (a, AA) 0.195% 99.805% 8 - Effect of repeated backcrossing on the frequency of homozygotes an Single locus A/a. (The proportion of AA also represents the proporti recurrent parent). id heterozygotes at a ion of genes from the266. 48.4.3. Selection for the Gene under Transfer insfer must be mail an efficient sele by z nt. Hf Iso be replaced by from the recurrent pare! will also be th ction in the backcross generation, ntained by The gene under tra re cllele Otherwise the gene 48.4.4. Extended Opportunit\ for Breakin: ndesirable Linkages y for Breaking Ul ig) . Average proportion of ‘The transferred gene and the genes tightly linked TABLE 18.1. A ig i genes from the re. Be ners Se here current parent in the ss generations. Therefore, u i ould pe roppertiity in each backcross different generations of generation for crossing over to occur between a backeross programme the gene being transferred and the genes tightly - “Average proportion linked with it, This is of great advantage when the Generation in parters) of pariés from linked genes are undesirable. However, when Ee ecurane paren genes are tightly linked (crossing over 0.1 per ae cent or less), large backcross populations would Fy 7 have to be grown to obtaine recombinant types. BC, as Often tightly linked genes may remain together BC; 87.5 up to BC 9. BC, 93.75 BC, 96.875 18.4.5. Selection for the Recurrent BCs 98.438 Parent Type BC, 99.218 Often a selection for the recurrent parent type is BC; 99.609 done during the backcross generations. This leads BC, 99.805 to a faster replacement of the genes of BC, 99,902 Ronrecurrent parent than that given in Table 18.1, Bw 99.951 Se 18.5. SELECTION OF PARENTS Itis clear that the backcross method of breeding changes the gen lotype of th y only that defect of the recurrent "ype of the recurrent parent only for Parent, which is aimed to the corrected 18.6. PROCEDURE OF BACKCROSS METHOD The plan of backcross methBackcross Method 267 18.6.1, Transfer of a Dominant Gene Let us suppose that a high yielding and widely adapted wheat variety A is susceptible to stem rust. Further, another variety B is resistant to stem rust, and the resistance to stem rust is dominant to susceptibility. A generalised scheme of the backcross programme for the transfer of rust resistance from variety B to variety A is given below (Fig. 18.2). Variety Bis the non-recurrent RST parent (rust resistant but Fa NONRECURRENT = RECURRENT agronomically undesirable). Variety Seri PARENTB = x PARENT A Ais the recurrent parent (rust An © susceptible but agronomically | desirable). eveand ia fir er (RECURRENT PARENT A) F, backcrossed with the recurrent parent, variety A. () Rust resistant plants similar mao we foes cin, ales sm YEAR a i /aponeecucemasaseie (iy Plants backcrossed with recurrent parent A. FOURTH ac, XG2, Rex rr(RECURRENT PARENT A) (i and (i) as in BC, FIFTH BC; wey, ArX rr(RECURRENT PARENT A) YEAR sictH ac, (irr), ReXer(RECURRENT PARENT A) —() and (i) as in BC, YEAR Wh SEVENTH BC, ey, RrX rr(RECURRENT PARENT A) (i) and (i) as in BC, YEAR () Rust resistant plants self- Pollinated. ‘Rr Seeds of selected plants ecu ag, re, (i) Fawestedseparatcy. YEAR: Ss @ al plant progenies fon CF || IIIIINIIIIILIN (iw Selection for rust resistance | and plant type of A. (i) Individual plant progenies TENT ecg, _|IIIIIIIIIIIIII © own. Co YEAR ( Homozygous progenies similar to recurrent parent A harvested and bulked. Replicated yield trial with the ov recurrent Parent as one of the YEAR 2 Wwe aes for distribution YEAR . . Fig. 18.2. generalised scheme for the transfer of a dominant gene for disease (rust) resistance through the backcross method in a selt-pollinated species. tr} denotes that pl : genotype are rejected. ey Plants with rr: Principles and Methoy | S | Plant, Breedin B. Generally, variety A should be used as 4, | f selfed plants. if any. 2 A. Since all the Fy plants wily 5, | stance is not necessary. | nts in (BC\) would be resistant and th, | Jants are selected and backcrossed i, :mblance to variety A as well, crossed to variety the identification ol re backcrossed tO variety tion for rust res! (BC,). Half of the pla tem rust. Rust resistant p) 1. Hybridization. Variety A is female parent. This would facilitate 2, Fy Generation. F plants a heterozygous for rust resistance, selec 3. First Backcross Generation remaining half would be susceptible to s i variety A. BC, plants resistant to rust may be selected for their rese! 4. BC,-BCs Generations. In each backeross generatio’s segregation would occur for rus, | resistance. Rust resistant plants are selected and backcrossed to the recurrent parent A. Selection for he plant type of variety A may be practised, particularly in BC, and BC3 generations. 5. BC Generation. On an average, BC, plants will have over 99% per cent genes from variety A. Rust resistant plants are selected and selfed; their seeds are harvested separately. i ke Bok: Generation: Individual plant progenies from the selfed seeds of the selected plants ae - stant plants simil 5 : ies cae conse plants similar to the plant type of variety A are selected and their selfed seeds arr 7. BCE; Gi ie i ‘Fy Generation. Individual plant progenies are grown, Progenies homozygous for rus Tesistance and similar to the plant type of variety in bulk. Several similar progenies are yp Aare harvested Ik. Severs 8 8. Yield Tests. Thi is i v Aas \e new variety is tested in a replicated yield trial along with the variety A a2 iet s check. Plant type, date of flo : wering, date of mat evaluated. Ordinaril aprihd barricade : eran Orla iy, the new variety would be identical to the vi Fae ete pele » generally not required and the variety may ee Sie performance. Detailed irectly released for cultivation. 18.6.2. Transfer of a Recessive Gene When rust (or an; i ¥y other disease) resista made one after the other. Aft bickomee ee After the first back “aes = » all the backcrosses cannot be 3. BC, Generation. Sit sista Current parent 1. Since rust resistance is Therefore, there is no test for rust resis noe nat geeessive, al ; st resistance, All the plants sf, tt th Plants bl Plants are ‘ill sceptib! Self-pors: ‘ill be rust suscepti 4. BC,F, Generation. Pla - Plants ar ‘ backcrossed with the recurrent patent. Selec with ust ad Parent. Selecti SPO variety A, the recurrent parent, mis done for the pan esistant pl ee Z plants are selecte 5. BC, Generation. There is 1. There is no rust resis recurrent parent A, and backcrossed with fee test. Plants are re sel nt Parent, Tected for their resemblance 10" t usually done for the plant type of variety A. est The plants are se 7. BCsF; Generation. Plants are inocu la : ted wit are selected and backcrossed to variety A. Selection forte Rust res or sis Plant type of ttt Plants resembling variety * variety Aj tY A is generally effectiVv®i packeross Method 269 RST NONRECURRENT RECURRENT ( Variety B is rust resistant YEAR PARENT B PARENT A Variety A is. susceptible but ea agronomically desirable Variety A and B are crossed: 8 isused as the female parent. ‘SECOND F, Rrx RR (RECURRENT (i) No rust resistance test. YEAR PARENT VARIETY A) (i) F; backcrossed to the recurrent parent. THIRD RA, Rr () No rust resistance test. YEAR me, \ (i) All piants are self-poliinated. (i) Rust resistance test. FOURTH BOF VARY Ser AR (VARIETY A) (i) Resistant plans selected and YEAR ran (il) backcrossed to variety A. FIFTH BC, ‘AX RA(VARIETY A) () No rust resistance test. YEAR / (i) Plants backcrossed to variety A. SIXTH BC; RR, Rr (0) No rust resistance test. YEAR \ All Plants selt-polinated. () Rustresistancettest. SEVENTH F, . Ce) rrX AR (VARIETY A) (i) Resistance plants selected oe a EIGHTH () No rust resistance test. YEAR BC eae ae (i) backcrossed to variety A. () No rust resistance test. NINTH sir BG, AR Rr (i) AllPlants self-poliinated. \ 0 a ad ad sistant selected an TENTH RGF; tm () Soited seeds are harvested YEAR 2 Separately. () Individual plant progenies ea ecsFs | (IIVIDMMMIIIIII 2 ‘ YEARS i Pr tesistant to rust similar to. variety A selected and composited. TWELFTH 4 Replicated yield test along with the YEAR variety A as the check. THIRTEENTH ‘Seed muttiptication for distribution YEAR begins. Fi 9 18.3. A generalised scheme for the transfer of a recessive gene for disease resistance (in this case rust resistance) through the backcross method in a self-pollinated crop. BC, Generation. There is no rust resistance test. Plants are backcrossed to variety A. ae Generation. There is no rust test. Plants are self-pollinated to raise F generation, rig BSF: Generation. Plants are subjected to rust epidemic. A rigid selection is done for rust “pely > and for the characteristics of variety A. Selfed seeds from the selected plants are harvested 7 estEE Plant Breeding Principles and Methog, 270 bjected to rust epiph: wn and subjec Piphyto ’ dual plant sare BO" f variety A. Seeds from ge 11, BC Generation, Indvidal plant poe Ta stics 0 mg rigid selection is done for resistance 10 ;xed to constitute the new variety. m igid : cous progenies are : similar rot resistant homogensohs DAT these steps are the same as in the Case OF transfer 12. Yield Tests and Seed Multiplicati dominant gene. 48.6.3. Number of Plants a ‘According to the above schemes, only a few (about yeneration for the .cter governed by a sing! F generati the transfer of a character g¢ i Tecan tainly have at least one plant with the gene for rust resistance. However, if the character is governgy cert nies would be required. A larger size g -e genes, a larger number of backcross progenies ‘ ie population is also desirable to permit an effective selection for the Lae felt of the ear parent, and to increase the probability of recombination beserean the sens ea paar el the ti ly 100 or moi ria jenes tightly linked with it. Therefore, more than 50, preferably 100 oF mé Each backeross ‘generation, In F, and F generations, the population size should be as large as possible 18.6.4, Selection for the Character Being Transferred A rigid selection for the character being transferred must be practised during the backcross and the, ‘generations, otherwise the character is very likely to be lost. It is, therefore, essential that the charac being transferred must have a high heritability. Although monogenic characters are the easiest transfer, the number of genes is not as important as the heritability of character under transfer. Iti desirable that the character should be easily identified either visually or through simple tests, including closely-linked DNA markers. The breeder should try to maintain the character in an intense fom through selection. Often some intensity would be lost due to modifying genes in the new geneic background. Therefore, the nonrecurrent parent should be chosen for a high intensity of the characte t0 be transferred. ever Necessary in Backcross Generations ) seeds are necessary in cach backer, Je gene. This population size Would alm, 18.6.5. Number of Backcrosses to be Made In the backcross method, it is essential that the genotype of recurrent parent is completely recover! except for the gene being transferred. Often the recurrent parent is likely to consist of several clos similar purelines. Therefore, a sufficient number of plants from the recurrent parent should be used (! the backcrosses. This would make sure that the new variety will have the same genet sition the recurrent parent, ea Generally, six backcrosses are sufficient to recover Selection for the characteristics of recurrent often have the effect of one or two additional the essential features of the recurrent pat Parent, Particularly in the early backcross generations, ™! backcrosses. Thus 4-5 backcrosses along with an effet!” years if a dependable phytotron was available, 18.6.6. Transfer of Leaf Rust Resistance to Malviya 12 Wh ic eat Malviya 12, a high yielding wheat variet i , y, became highly susceptibl wit? : ‘Pile to stem and leaf rusts we) * the three rusts under artificial inoculatio’ (Malviya 12 x Sparrow), 100 Plants were grown; all of the™= packeross Method 271 a backcrossed to Malviya 12. In BC,, more than 500 plants were grown. Epidemic condition was created by planting infector rows of Agra Local, a wheat variety susceptible to all the known races of stem, leaf and stripe rusts. The inoculum consisted of the naturally occurring races of leaf and stem rusts; no artificial inoculation was done. One hundred rust resistant plants were selected and backcrossed to Malviya 12, In BC, and BC; generations, the plants were handled in the same way as in the BC;. In BC, all the rust resistant plants (about 300) were selfed, and BC,F, was raised. Rust resistant plants were selected and selfed in BC,F>, and individual plant progenies were grown in BC,F; 50 progenies homozygous for rust resistance were selected and their seeds were bulked to constitute the rust resistant Malviya 12. Yield trials revealed a slight (about 4%) improvement in yielding ability of the new variety over the recurrent parent Malviya 12. During the backcross programme, the sole criterion of selection was rust resistance. One hundred rust resistant plants were selected in each generation to ensure that resistance to both stem and leaf rusts was selected for, and that at least some of the resistant plants were not escapes. 18.6.7. Transfer of Quantitative Characters Quantitative characters with moderate to high heritability, such as, grain size, plant height, flowering or maturity time, etc., are suitable for transfer through the backcross method. Invariably, some intensity of the character will be lost during the transfer. Therefore, the nonrecurrent parent must have the character in a more intense form than it is desired in the new variety. After each backcross, the progeny are selfed and F2 of 500-1000 plants is grown. A rigid selection for the character being trasferred is practised in each F2 generation. At the end of backcross programme, the F, is handled according to the pedigree method. Several progenies homozygous for the character being transferred and similar to the recurrent parent in other characteristics are selected and mixed to make up the new variety. An example of such a transfer is provided by the medium-grained rice variety Calady. Calady was developed through four backcrosses of the F hybrid betwcen Lady Wright (a long-grained variety) and Coloro (a short-grained variety) to Coloro. After each backcross, F, was grown and a rigid selection was practised for grain size, 18.7. TRANSFER OF TWO OR MORE CHARACTERS INTO A SINGLE RECURRENT PARENT When two or more characters are to be transferred into the same variety, one of the following three approaches may be used to achieve this objective. 18.7.1, Simultaneous Transfer Genes for the different characteristics may be transferred simultaneously in the same backcross Programme. The characters to be transferred are brought together into the hybird by successively crossing each of the nonrecurrent parents to the recurrent parent or the hybird thus produced. But in Such a case, a larger backcross population would be needed than in the case of transfer creer Further, the breeding programme may be delayed because the conditions neces: Selection of all the characters may not occur every year. Sometimes, the two ger linked. In such a case, the transfer becomes very easy, Benes under transfer may and selection for only one "ecessary. Some examples of such a favourable linkage are between the dSro4 tro genes L724 and $725, and L726 and $731. easter of a single sary for the 18.7.2. Stepwise Transfer Ih a Approach, the recurrent parent is first improved for one charater. The i : impro used as the recurrent parent in a fres! Proved recurrent parent fh backcross programme for the transfer of the second >,Plant Breeding * © hon, 272 sa : they are transferred one at a time in a Step, J, tes 8 are to be transfert neler of two oF more chraracters. Jn longer time for the t 48.7.3. Simultaneous but Separate Transfers ¢ same recurrent pare! character. If additional characters fashion. This approach takes muc! nt in simultaneous but separate backer, arent resulting from the different backeryy characters being transferred arg then hod. This approach appears to be iy 4 esti Each character is transferred into ¥ programmes. The improved versions of the recurrent patel programmes are then crossed together. Homozygous lines f0 Selected from the segregating generations using the pedigree met most suitable of the three strategies. MODIFICATIONS OF THE BACKCROSS METHOD ways to suit the needs of various breeders The backcross method may be modified in variot Following are the three common modifications of the backcross method. 18.8.1, Production of F, and F, F, and Fy generations are produced after the first and the third backcrosses. A rigid selection for thy character being transferred and for the characteristics of the recurrent parent is done in the F, and f, | generations. In the backcrosses progenies, selection need not be done either for the character being transferred or for the characteristics of the recurrent parent. The fourth, fifth and sixth backcross a | made in succession. For the sixth backcross, a relatively larger number of plants from the back-cros, Progeny are used. This method may be used for the transfer of both dominant and recessive genes. lis believed that an effective selection in F; and #3 generations is equivalent to one or two additional backcrosses. 18.8.2. Use of Different Recurrent Parents ‘1-2 times to the recurrent Progeny are handled according to the pedigree method. This approach is useful why pews seful when one of the pa Parent. Subsequently, the backcross Tistics but the no} nre: Rh pa as the recurrent pee ent Parent is not undesirlBackcross Method 273 ae 18.9. APPLICATIONS OF BACKCROSS METHOD TO CROSS-POLLINATEN CROPS The backcross method is equally applicable to cross-pollinated crops. The method used for eress- pollinated crops is essentially the same as that applied to self-putlinated crops. ‘The only differer ~ is that in cross-pollinated crops a large number of plants (100-3vu) som the recurrent parent must be used in each backcross. This is necessary so that the new variety has the sane genetic constitution as the recurrent parent. For example, wilt resistance was transferred into alfalfa variety California Common from the variety Turkestan, Two hundred plants of California Common were used for each backcross. The new variety Caliverde is exactly like California Common, except for its wilt resistance. 18.10. MERITS OF BACKCROSS METHOD 1, The genotype of new variety is nearly identical with that of the recurrent parent, except for the genes transferred. Thus the outcome of a backcross programme is known beforehand and it can be reproduced any time in the future, 2. It is not necessary to test the variety developed by the backcross method in extensive yield tests because the performance of the recurrent parent is already known. This may save up to 5 years’ time and a considerable expense. 3. The backcross programme is not dependent upon the environment, except for that needed for the selection of the character under transfer. Therefore, off-season nurseries and green-houses can be used to grow 2-3 generations each year. This would drastically reduce the time required for developing the new variety. 4. Much smaller populations are needed in the backcross method than in the case of pedigree method. 5. Defects, such as susceptibility to disease, of a well-adapted variety can be removed without affecting its performance and adaptability. Such a variety is often preferred by the farmers and the industries to an entirely new variety because they know the recurrent variety well. 6. This is the only method for interspecific gene transfers, and for the transfer of cytoplasm. 7. Itmay be modified so that transgressive segregation may occur for quantitative characters. 18.11, DEMERITS OF BACKCROSS METHOD 1. The new variety generally cannot be superior to the recurrent parent, except for the character that is transferred. 2. Undersirable genes closely linked with the gene being transferred may also be transmitted to the new variety. 3, Hybridization has to be done for each backcross. This is often difficult, ime taking and costly, 4, By the time the backcross programme improves it, the recurrent parent may have been replaced by other varieties superior in yielding ability and other characteristics. 18.12, ACHIEVEMENTS The backcross method has been widely used in crop improvement. It has been used for transferrin, genes and chromosomes from one variety to another and from related species, and for the transfer cf cytoplasm, Backcross method is the only breeding method suitable for the transfer of genes and chromosomes from one species to another, There are many examples of interspecific transfer of using the backcross method. ‘Transfer’ was the first commercial wheat variety in which rust res anse Was transferred from a related species. st resistnce Two cotton varieties 170-Co2 and 134-Co2M, were develo} ; d using the bac! hirsutum veriety Dharwar American 2-6-5 was crossed with G. crborewe ge cree method. G. arboreum variety Gaorani 6. The F,ova Plant Breeding : Principles and Methog, s. Which gave rise to tetraploid plants (perhaps as a result Of backer, te the G_ hirsutum parent), Progeny from the F, were backcrossed to G. hirsutunt varieties Dharwa, frst hackeross), Comboxtia Co? (second backcross) and Meade (third backcross). From y, American (first backcross), Combodia a 12 (seer ‘These varieties bad staple length a _ backcross progeny, 170-Co2 and 134-Co2M were selected. Thes eae I 1 (about 2.7 em) and were widely cultivated in Gujarat under irrigated and rainfe S. ‘The backcross method has been extensively used for the transfer of disease resistance to popular widely atapted varieties, This is illustrated by the popular wheat (7. aestivum) variety Kalayan Sona, which became susceptible to leaf rust. Rust resistance has been transferred to Kalyan Sona from severg diverse sources, ¢,g., Robin, KI, Bluctrird, Tobari, Frecor, HS19, etc., using the backcross method ‘Three multiline varieties, viz, KSML3, MLKSI1 and KML7406, have been released for cultivation ag a result of the above (Table 18.2), Another example is provided by the hybird bajra varieties in India The male sterile line Tif23A (the female parent in the hybrids) was highly susceptible to downy mildew leading to a high susceptibility of the hybirds. Tift23A was used in backcross programmes with resistant lines trom India and Africa to develop downy mildew resistant male sterile lines, such as MSS21, MSS41A, MSS70A, ete., which are now being used for producing downy mildew resistant hybinds, was highly sterile: it set few se ia v TABLE 18.2. Indian wheat varieties developed through the backcross method Voriety evolved Recurrent parent Non-recurrent parent Pureline Verieties Kharchia 65 Kharchia local EG953 NPSS2 NP761 EG9S1 NIS439 NP7L0 REPM80 rwe72 Kalyan Sona. E6056, HUW234 HUWI2 CPAN1666 (One backcross) K8027 K8S2 HD1696 W204 C306 * Multiline Varieties MLKS11 (8 components) Kalyan Sona Exotic lines resi s is resistant to leaf rust KML7406 Kalyan Sona Exotic lines resistant to leaf rust eens Kalyan Sona Sonalika Exotic lines resistant to leaf rust Six parents ** Leaf rust resistance due to Lr24 (character that was transferred), —_—_ #* Released for Punjab state Characters other than disease resistance have been transferred examples of this types are available in cotton. G. herbaceum varie, Kalyan are some of the cotton varieties developed by the backcross Local, a mixture of Goghari and Broach Desi varieties of G, Goghari A-26 and Broach Desi 8 (BD8), were selected. Variety Bi spinning value, but it had a low ginning outturn (34%). BD8 w; backcross programme with Goghari A-26, which had a high gin ourtturn was transferred from Goghari A-26, the nonrecurrent developed. Vijay was further improved through the backcross meth, maturity were transferred from 1027A L-F (nontecurrent parent), above backcross programme. herba fe vaccum ‘wo pureline varieties. eo esistant and had a high by a backcross method. Good saa Vijapla, Vijay, Digvijay and |. From the variety Broach re ning outtum (42 qamrent Parent in a 7%), High ginning ‘aricty Vijay was ved fibre length and early ‘Svijay resulted from the Parent, and a 1od; improved Variety D;kcross Method Dae eo eee 18.13, COMPARISON BETWEEN BACKCROSS AND PEDIGREE METHODS The wo muathade 7 only cites in their procedure and the system of mating for raising the generations after Fy; they also differ in their end products as well. These differences are summarised in Table 18.3. TABLE 18.3. Comparison between backcross and pedigree methods pollinate ‘The new variety developed by this method is different from the parents in agronomic and other characteristics ‘The new variety has to be extensively tested before release ‘The method aims at improving the yielding ability and. other characteristics of the variety It is useful in improving both qualitative and quantitative characters It is not suitable for gene transfer from related species Backcross method the recurrent parent ‘The new variety is identical with the recurrent parent, except for the character under transfer Usually extensive testing is not necessary before release ‘The method aims at improving specific defects of a well adapted, popular variety It is useful for the transfer of both quantitative and qualitative characters provided they have high heritability It is the only useful method for gene transfers from related species and for producing addition and substitution lines Hybirdization with the recurrent parent is necessary for producing every backcross generation The backcross generations are small and usually consist of 20-100 plants in each generation ‘The procedures for the transfer of dominant and recessive genes are different and for producing substitution or addition lines (ed to the production of the Fi generations ‘The Fr and the subsequent generations are much larger than those in the backcross method ‘The procedure is the same for both dominant and recessive genes Questions 1. Define the following: Recurrent parent, Nonrecurrent parent, Backcross, BC,F;, BC, By and 8. 2. List the requirements for a backcross programme. Describe the procedure of backcross method suitable for the transfer of a dominant gene for disease resistance. 3. Describe the procedure of backcross method for the transfer of a recessive gene. Discuss the merits and demerits of the backcross method of breeding. ‘4. Describe the applications of the backcross method and Its achievements. 5. Write short notes on the following: (/) Transfer of quantitative traits through backcross method, (W) Application of backcross method to cross-pollinated species. (iii) Selection in backcross. (i) Modifications of backcross method, (¥) Number of plants necessary in a backcross geeaanrand (wv) Number of backcrosses required In gene transfer programmes, Suggested Further Readings SaaS ‘Alard, R.W. 1960, Principles of Plant Breeding. John Wiley and Sons, Inc., New York. riggs, F.N, 1959, Backerossing—its development and present application. Proc. ist International Wheat Genetice Symposium, pp 8-9. 1 Nand Allard, R.W. 1953. The curent sts of backcross method of plant breeding. Agron 3. 4S: 131- Raddy, avs, ,LV.S, and Comstock, R.E. 1976, Simulation of the backcross breeding method. I, Effects of ‘and gene number on fixation of desired alleles. Crop Sc. 16: 825-830, erteabity _