Multiple factor

It is quite natural that small differences exist among individuals of similar genotype due to the effect of environment on genotype. On the other hand, there are some heritable differences also exist with continuous variation. Most of the economical traits show continuous variation and they are measurable or quantifiable. Quantitative characters Quantitative characters or traits which show continuous variation and governed by a large number of genes called multiple genes or multiple factors or polymeric genes or polygenes. Their inheritance follows same mendelian principles.

Qualitative characters Qualitative characters show discontinuous variation and are governed by one or two major genes or oligognes.

Multiple factor Hypothesis (Nilson Ehle)

Nilson-Ehle studied Kernel colour in wheat concluded that is a quantitative character He crossed true breeding red kernel wheat (RR) with true breeding white (rr) and the F1 was red (Rr) and the F2 segregated for red and white in 3:1 ratio indicating the dominance of red over white. However, careful examination indicated the variation in red color among the red color progenies F1 red was not as intense as one of the parents In F2 he could observe two grades of red ie., one was red as that of one of its parent, two were higher red as that of F1 individuals. In some crosses, a ratio of 15 red : 1 white was found in F2 indicating that there are two pairs of genes for red colour that either or both of these can produce red kernels.

o Finally he observed different shades of red in F2 for red kernel types. The F2 showed red shades and white as follows; Dark red :1 Medium dark red : 4 Medium red :6 Light red :4 White :1 Total : 16 It was concluded two duplicate dominant alleles R1 and R2 cumulatively decide the intensity of red colour and both R1 and R2 are in completely dominant over white. The high intensity of red colour depends on the number.

The F2 ratio in wheat Genotype R1R1 R2R2 R1R1 R2r2 R1r1 R2R2 R1r1 R2r2 R1R1 r2r2 r1r1 R2R2 R1r1 r2r2 r1r1 R2r2 r1r2 r2r2 Genotypic ratio 1 2 2 4 1 1 2 2 1
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Phenotype Dark red Medium dark red Medium dark red Medium red Medium red Medium red light red light red white

 Hence, if two parents differ for the two genes the segregation was 1:4:6:4:1 provided both R1 and R2 contribute equally to the colour. o If three genes are involved in F2 segregation showed 1:6:15:20:15:6:1 for red shades and 1 for white. Thus, Nilson-Ehles multiple factor states that Multiple factor hypothesis (Nilson - Ehle) i) for a given quantitative trait there could be several genes, which were independent in their segregation, but had cumulative effect on phenotype ii) Dominance is usually incomplete iii) Each gene contributes something to the strength of expression of character whereas its recessive allele does not of genes present dominance gene. Multiple factor Quantitative inheritance The alleles that contribute for the trait are called "contributing alleles",

If four genes are determining the kernel color then,

R1R1 R2 R2 dark red

r1r1 r2r2 white

F1 R1r1 R2r2 Medium red

 R1R2

R1R2

R1r2

r1R2

r1r2

R1R2 R2R2
(dark red)

R1R1 R2r2
(Medium dark red)

R1r1 R2R2
medium dark red

R1r1 R2r2
light red

R1r2 r 1R 2 r 1r 2

R1R1 R2r2
medium dark red

R1R1 r2r2
light red

R1r1 R2r2
light red

R 1r 1 r 2r 2
very light red

R1r1 R2R2
medium dark red

R1r1 R2r2
light red

r1r1 R2R2
light red

r1r1 R2r2
very light red

R1r1 R2r2
light red

R1r1 r2r2
very light red

r1r1 R2r2
very light red

r 1 r 1 r 2r 2 white

`Dark red is due to the presence of four contributing genes medium dark is due to three genes and medium red is due to two contributing genes and light red is due to one contributing gene

One gene segregation in quantitative

Two gene segregation

If N genes are heterozygous in the F1 hybrid, there will be: 2N +1 phenotypic classes in the F2 (0, 1, 2, 3, etc. up to 2 N contributing alleles).
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 The fraction of individuals falling into each of the extreme classes (0 or 2N contributing alleles) will be 1/4th to the N power (1/4)

Transgressive segregation
The appearance in f2 of individuals with a higher or lower intensity of character than those parent in the parents involved in the cross is termed as transgressive segregation. Clearly transgressive segregants transgress or surpass the parental limits or a quantitative trait, and they are consequences of segregation. Eg: Inheritance of kernel colour in wheat. The nature of transgressive segregation in plants A remarkable diversity of traits have been shown to exhibit transgression in plants. The bulk of these are morphological traits (65%), whereas the remainder are fairly evenly divided among trait categories such as fecundity, the biochemical composition of organs and tissues, physiology, life history, and tolerances to various biotic and abiotic factors (Appendix 1). This latter category may be most important to the success of hybrids, because transgressive segregation for ecological tolerances seems most likely to facilitate niche divergence. Examples of transgressive segregation for abiotic factors include increased cold, salt, drought, heat, and heavy metal tolerances. Likewise, transgressive phenotypes for resistance to pathogens and herbivores are often reported. Of course, we recognize that expanded ecological tolerances may be a secondary consequence of transgressive changes in
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morphology, life history, physiology, or biochemical composition.

The appearance of individuals in F2 with very higher or lower intensity of expression than their both parents is known as transgressive segregation and such individuals are called transgressive segregants. Transgressive segregants are produced when i) the two parents involved in a cross which have positive alleles of different genes affecting a quantitative characters. segregation for these genes produces the two extreme homozygotes in F2 which transgress the parental limits for the character

Polygenes
Thus, the studies of Nilsson-Ehle demonstrated the existence of several genes with small cuimulative effects governing a single trait; such genes are commonly known as poly genes(sometimes minor genes ). In contrast genes having a large effect on the characters they govern are called oligigenes or major genes. Characters governed by polygenes (multiple factor ) but showing discrete variations are called quasi quantitative characters.

The differences between qualitative and quantitative traits

effect of individual contributing genes is not distinguishable continuous variation quantative traits are measured on a continuous scale (mean, variance) with statistical method) 6. Influence of environment More influence of environment is very less 7. Dominance is complete each dominant gene has a in mendelian ratio quantitative effect 8. Lesser role of modifying more role of modifying genes genes