a . In case of oligogenic characters, the gene action (dominance and epistasis). a 8, Incase of polygenic characters, metric measurements like size, weight, duration, strength, etc, are possible, whereas in case of oligogenic characters only the counting of plants with regard to various kinds like colour and shape is possible. Thus, metric measurement is not possible jn case of oligogenic characters. 9. Transgressive segregants are only possible from the crosses between two parents with mean values for a polygenic character. Such segregants are not possible in case of qualitative or oligogenic traits 10, The transmission of polygenic characters is generally low because of high amount of environmental variation. On the other hand, oligogenic characters exhibit high transmission because there is little difference between the genotype and phenotype of such character. Thus, polygenic characters differ from oligogenic ones in several aspects (Table 19.1). TABLE 19.1 Differences between polygenic and oligogenic traits | Polygenic Traits Oligogenic Traits 1. Governed by several genes. Governed by few genes. 2. Effect of each gene is not detectable. Effect of each gene is detectable. 3. Usually governed by additive genes. Governed by non-additive genes. 4. Variation is continuous. | Variation is discontinuous. 5. Separation into different classes is not possible. | Separation into different classes is possible. 6. Highly influenced by environmental factors. Little influenced by environmental factors. 7. Statistical analysis is based on mean, variances | Statistical analysis is based on frequencies or) |___and covariances. ratios. In plant breeding both types of characters showing qualitative and quantitative inheritance have equal economic importance. SIMILARITIES BETWEEN OLIGOGENIC AND POLYGENIC TRAITS East (1916) demonstrated that polygenic characters were perfectly in agreement with Mendelian segregation and later on Fisher (1918) and Wright (1921, 1935) provided a mathematical basis for the genetic interpretation of such characters. The quantitative characters do not differ in any essential feature from the qualitative characters (Falconer, 1960, 1981 ; Mather, 1949 : Mather and Links, 1971), as discussed below : 1, Both quantitative and qualitative characters are governed by genes, the former is controlled by polygenes or minor genes and the latter by oligogenes or major genes Both major as well as minor genes are located on the chromosome in the nucleus, 3. The polygenic traits controlling continuous variation exhibit segregation like major genes controlling discontinuous Mendelian variation, 4, Polygenic characters show variable expression which environmental effects. Quali ‘tive -haracters also exhibit vz degree than polygenic traits due to non-genetic causes i on in expression but to a lesser 5. The reciprocal crosses for both types of traits exhibit close agreement in expression of genes.exhibit linkage like oligogenes. Many i we ma od F é cases of linkage between major genes and controlling continuous variation have been reported. = ; ; , quantitative genetics or biometrical genetics is se eaeratianrata ieee int Mendelian principles of heredity, stan a ie a POLYGENIC TRAITS pic 1 of analysis of quantitative inheritance differs trom that of qualitative inheritance in some is as given below 1. Ttrequires various measurements of characters like weight, length, width, height, duration, ete.. rather than classification of individuals into groups based on colour or shape. Ds Observations are recorded on several individuals and the mean values are used for genetical studies. Segregation into distinct classes in F, generation is not obtained in the inheritance of quantitative characters. The segregants exhibit continuous range of variation from one extreme (low) to other (high) for such traits. 3. The inheritance is studied with the help of mean, variances and covariances. These estimates can be worked out from data recorded in replicated experiment. 4, Fisher (1918) was the pioneer worker to interpret the quantitative characters in terms of Mendelian genetics. Now several biometrical techniques are available for the genetic analysis of quantitative characters. The science which deals with the genetic interpretations of quantitative characters has got separate entity as quantitative genetics or biometrical genetics. ASSUMPTIONS OF POLYGENIC INHERITANCE Polygenic inheritance is based on several assumptions. The six important assumptions are given below : a contributing alleles and those which do not contribute to continuous vari ‘contributing alicles. Some scientists refer to these as effective and non-effective allele: 1. Each of the contributing genes involved in the expression of a character produces an equal effect. 2, Each contributing allele has either cumulative or additive effect in the expression of a character. 3. The genes involved in the expression of characters have lack of dominance. They show intermediate expression between two parents is among genes at a different loci. 4, There is no epist 5. The linkage is in equilibrium, means there is no linkage. 6. The environmental effects are absent or may be ignored. However, seldom fulfilled. There are two types of alleles or genes in the polygenic inheritance, viz., 1. contributing alleles and non-contributing alleles. Those alleles which contribute to continuous variation are known as tion are referred to as non- respectively. ast three assumptions areitance has been reported for various characters ‘examples include kernel colour in wheat, corolla length maize. These are briefly described as follows : 7 Kernel Colour in Wheat . oa Nilsson Ehle (1908) studied the inheritance of kernel colour in wheat. He found that seed or kerne} colour in wheat is governed by one, two and three gene pairs, because in the crosses between red and white kernel varieties, he observed that the F, was intermediate between the parental values and in F, hhe observed 3 : 1, 15: | and 63: 1 ratios of red and white seeds in different crosses. The last two ratios indicated that there was duplicate gene interaction, However, indepth study of coloured seeds revealed that there were different grades or shades of colour within the red coloured seeds. The red seeds of 15 * 1 ratio could be easily divided into four classes on the basis of shade of colour, viz., dark red, medium dark red, medium red and light red. These colours were observed in the ratio of 1; 4:6: 4: 1. This Suggested that the seed colour in wheat is controlled by genes which show lack of dominance and have small cumulative effects. Here, two types of alleles are involved in the expression of character. Those which contribute to continuous variation and those which do not contribute. The first category of alleles is called effective and second as non effective. Assume that red seed colour is controlled by two genes R, and R; and, white seed colour by r, and r,. From the cross between dark red and white seed parents, Nilsson Ehle observed the following results (Fig. 19.1) : Parents Dark red. White Genotypes RRRR x Htefe 4 F, RyRy, Medium Red R,Re itp Re iy F, RiP, RiR,R-Rp RR Ror yrs RoRe Furst {OR} IMDR} [MOR] (MR) Rite RRR, RiRitote RirRors Bitters {MOR} (MRI IMR} [LR] Re RirsRiaRe RitRote nnR AR Tt Pate [MDR) [MR] {MR] (LR) Ne RyRere Rititofe Rory Thyfore [MR] {LR} (LR) DR = Dark Red, MDR = Medium Dark Red, MR = Medium Red, LR = Light Red and W = White Fig. 19.1. Inheritance of kernel colour in wheatis 0 divid the genetic variance into additive, dominance and epistatic ents. These are briefly discussed below (Mather and Jinks, 1971). fixable and, therefore, selection for traits governed yariance is important for the following major reaso} 1. Itis required for estimation of herit ability proportionate to narrow sense heritability. 2, Ilsa pre-requisite for selection because this is the only variance which responds to selection. 3. Breeding value of an individual is measured directly by the additive gene effects, The general Combining ability (gca) effect of a parent is measure of additive gene effects, 4. Additive genetic variance gets depleted proportionate to the improvement made by selection. >. In natural plant breeding populations, additive variance is the predominant one closely followed by dominance variance. DOMINANCE VARIANCE It arises due to the deviation from the additive scheme of gene action resulting from intra-allelic Interaction i.¢., interaction between alleles of the same gene or same locus. It is due to thé deviation of heterozygote (Aa) from the average of two homozygotes (AA and aa). Such genes show incomplete, complete or over-dominance. The dominance variance is associated with heterozygosity and, therefore, itis expected to be maximum in cross-pollinating crops and minimum in seif-pollinating species Dominance variance is not fixable and, therefore, selection for traits controlled by such variance is not effective. Heterosis breeding may be rewarding in such situation, Dominance variance differs from in narrow sense and response to selection is directly additive variance in several ways (Table 19.2). TABLE 19.2 Differences between additive and dominance variance Additive variance | ____ Dominance variance | It refers to difference between homozygotes | it refers to deviation of Aa from the mean of AA (AA/aa). and aa. | 2 Genes show lack of dominance. Genes show incomplete, complete or over- dominange, 3 Associated with homozygosity and,is more in| Associated with heterozygosity and is more in nin eeders | outbreeders, 4. Itis fixable, | {tis non-fixable. Selection is very effective as it is fivable. | Selection is ineffective as it non-fixable the chief cause of trans ive segregation. | It is the chief cause of heterosis or hybrid vigour.@ Additive x Additive In this case both the interacting loci exhibit lack of dominance individually. It is denoted as A x A and is fixable. (i) Additive x Dominance It refers to interaction between two or more loci, one exhibiting lack of dominance and the other dominance individually. It is denoted as A x D and is non-fixable. Gif) Dominance x Dominance In this type of epistasis both the interacting loci e D x D and is non-fixable. The first type of epistasis is fixable and, therefore, selection is effective for traits govemed by such variance. The last two types of epistatic variances are unfixable and, therefore, heterosis breeding may be rewarding for traits exhibiting such variance. In natural plant breeding populations, epistatic variance has the lowest magnitude. Epistatic variance differs in many aspects from dominance variance (Table 19.4). Wright (1935) suggested the partitioning of genetic variance into two components, viz., additive and non-additive (dominance and epistatic components), of which only the additive component Contributes to genetic advance under selection. Mather (1949) divided the phenotypic variance into three components, namely, 1. heritable fixable (additive variance), heritable non-fixable (dominance and epistatic components), and 2. non-heritable non-fixable (Environmental fraction). In fact, the ibit dominance individually. It is represented as