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GENETIC AND BREEDING INVESTIGATIONS IN TOMATO (Lycopersicon esculentum Mill.)

Thesis submitted to the University of Agricultural Sciences, Dharwad in partial fulfillment of the requirements for the Degree of

MASTER OF SCIENCE (AGRICULTURE)

In

GENETICS AND PLANT BREEDING

By PURNANAND G. KULKARNI

DEPARTMENT OF GENETICS AND PLANT BREEDING COLLEGE OF AGRICULTURE, DHARWAD UNIVERSITY OF AGRICULTURAL SCIENCES, DHARWAD - 580 005 AUGUST, 2006

ADVISORY COMMITTEE

DHARWAD AUGUST, 2006 Approved by : Chairman :

(O. SRIDEVI) MAJOR ADVISOR _________________________ (O. SRIDEVI)

Members : 1. ________________________ (P. M. SALIMATH)

2. ________________________ (M. S. PATIL)

3. ________________________ (R. M. HOSAMANI)

LIST OF ABBREVIATIONS

F1s BIP's Vg Vp GCV PCV h²(bs) GA

= MHTM256 and S-4-14 = Biparnetal progenies = Genotypic variance = Phenotypic variance = Genotypic coefficient of variation = Phenotypic coefficient of variation = Broadsense heritability = Genetic advance

GA (as % mean) = Genetic advance as per cent mean

CONTENTS

Chapter No.

Title

I

INTRODUCTION

II

REVIEW OF LITERATURE

III

MATERIAL AND METHODS

IV

EXPERIMENTAL RESULTS

V

DISCUSSION

VI

SUMMARY

VII

REFERENCES

VIII

ABSTRACT

LIST OF TABLES

Table No. 1.

Title

Review of literature on comparison of variability, heritability and genetic advance for various characters in BIPs and selfed population in wheat Review of literature on comparison of variability, heritability and genetic advance for various character in BIPs and selfed populations in safflower Review of literature on correlation studies in BIPs and selfed population in wheat Heterosis for different traits in tomato as reported by various authors Review of literature on combining ability in tomato Salient features of parents used in diallel mating Mean performance of F3 population, check and BIP populations in respect of nine characters in tomato Range for nine traits in F3 population, check and BIP population in tomato Estimate of genetic parameters for nine characters in biparental and F3 populations of tomato Nature of association among nine traits in biparental (BIP) Population M of tomato Nature of association among nine traits in biparental (BIP) Population S of tomato Nature of association among nine traits in biparental (BIP) Population M/S of tomato Nature of association among nine traits in biparental (BIP) Population S/M of tomato

2.

3.

4.

5. 6. 7.

8.

9.

10.

11.

12.

13.

Contd...

Table No. 14.

Title Nature of association among yield and different characters in F3 Population of tomato Path coefficient analysis of different characters towards fruit yield per plant in BIP Population M of tomato Path coefficient analysis of different characters towards fruit yield per plant in BIP Population S of tomato Path coefficient analysis of different characters towards fruit yield per plant in BIP Population M/S of tomato Path coefficient analysis of different characters towards fruit yield per plant in BIP Population S/M of tomato Path coefficient analysis of different characters towards fruit yield per plant in F3 population of tomato Frequency distribution of superior segregants for fruit yield in different segregating populations of tomato Analysis of variance in respect of 18 characters in a set of 7x7 half diallel crosses in tomato Per se performance, nature and magnitude of heterosis for plant height and primary branches in tomato Per se performance, nature and magnitude of heterosis for flowers per truss and fruits per truss in tomato Per se performance, nature and magnitude of heterosis for fruits per plant and average fruit weight in tomato Per se performance, nature and magnitude of heterosis for locules per fruit and fruit shape index in tomato

15.

16.

17.

18.

19.

20.

21.

22.

23.

24.

25.

Contd...

Table No. 26.

Title Per se performance, nature and magnitude of heterosis for pericarp thickness and ascorbic acid in tomato Per se performance, nature and magnitude of heterosis for total acidity and reducing sugar in tomato Per se performance, nature and magnitude of heterosis for nonreducing sugar and total sugar in tomato Per se performance, nature and magnitude of heterosis for juice recovery percentage and pulp content in tomato Per se performance, nature and magnitude of heterosis for fruit yield and total soluble solids in tomato Analysis of variance for combining ability in tomato General combining ability effects of seven parents for eighteen quantitative and quality traits in tomato Specific combining ability effects for eighteen quantitative and quality characters in F1 hybrids of tomato Per se performance of individual superior segregants for nine characters in different BIP and F3 populations in tomato Relative performance of top 3 hybrids with respect to per se value and check for 18 quantitative and quality traits in tomato

27.

28.

29.

30.

31. 32.

33.

34.

35.

LIST OF FIGURES

Figure No.

Title

1.

Estimate of GCV and PCV (%) for fruit yield in biparental and F3 populations of tomato

2.

Estimate of GCV and PCV (%) for fruits per plant in biparental and F3 populations of tomato

3.

Per cent heterosis for top three hybrids for yield over commercial check

LIST OF PLATES

Plate No.

Title

1.

Fruit shape and size variation observed in populations developed through intra mating

2.

Fruit shape and size variation observed in populations developed through selfing

3.

Fruit shape and size variation observed in populations developed through intermating

4.

Superior segregant for fruit yield observed in MHTM-256 X S-414 population (3.34 kg/plant)

5.

Fruit features of top heterotic hybrids

6.

Fruit features of hybrids involving Local type as one of the parents

I. INTRODUCTION

Tomato (Lycopersicon esculentum Mill.), is one of the most important Solanaceous vegetable crops of Peru-Ecuador origin (Rick, 1969). It is an important source of minerals and vitamins (Vitamin A and C). Because of high nutritional quality tomatoes are being used directly as raw vegetable in sandwiches, salads etc. Several processed products like paste, puree, soup, juices, ketchup, drinks, whole peeled tomatoes etc., are prepared on large scale and enjoy high acceptance as food ingredients. Thus, today it is one of the important raw materials for multimillion food industries. The present Indian annual production of 7.6 million tonnes from an area of 5.4 lack hectares (Anon., 2005) is not sufficient to meet the requirement of fresh market and processing industries. Karnataka is one of the important tomato growing states covering an area of 0.37 lack hectares with a production of 1.02 million tonnes (Anon., 2004). Conventional breeding methods such as pedigree, bulk and backcross breeding with some modifications have been principal procedures followed in the improvement of self pollinated crops. Such procedures though significant and productive in their own right impose restriction on the chance of better recombination because of larger linkage blocks associated with the weakness of causing rapid homozygosity and low genetic variability (Clegg et al., 1972). Further, negative correlations among yield components and high genotype x environmental interactions prevent full exploitation of genetic variability for characters like yield. If these propositions are to be accepted, we must reevaluate our breeding procedures. Presence of linkage blocks and inverse relations among the correlated characters are more common in tomato as it is a self pollinated crop. Under such circumstances, the biparental mating in an appropriate segregating material/population is effective in breaking larger linkage blocks and provides more chances of recombination than the selfing series (Gill, 1987). It is a useful system of mating for rapid generation of variability and may appropriately be applied where lack of desired variation is the immediate bottleneck in the breeding programmes. In any crop, yield is a complex character influenced by many of its contributing characters, and is controlled by polygenes and their interaction with environmental factors. So, an understanding of inheritance and study of association between fruit yield and its component traits is necessary for planning effective selection programme in identifying high yielding genotypes. However, the inheritance of quantitative traits is often influenced by variation in other characters which may be due to pleiotrophy or genetic linkage (Hanson et al., 1956). Hence knowledge of association between yield and its attributes is necessary in selecting suitable genotypes. Conventional methods of handling the segregating populations like pedigree method or bulk method do not provide any opportunity for the reshuffling of genes. Hence, any unfavourable association observed in an early segregating generations like in F2 are likely persist through the filial generations. However breeders can alter such association by resorting to approaches like biparental mating in the segregating populations. Such approaches have been tried and positive results have been reported in some crops like wheat, safflower etc. (Nanda et al., 1990). However no such report is available in tomato. Further the biparental mating of the selected plants of F2s of population is usual process. But mating between plants which are selected in F2s of one population and F2s of other population is a deviation of standard BIP mating, may be tried to increase the opportunity for extra recombination and consequently release higher variability. To meet the ever increasing demand for this vegetable, there is a need for development of hybrids and varieties with improvement in yield, quality, resistance to pest and diseases. Heterosis breeding as a tool for genetic improvement in tomato has been advocated by several workers ever since the phenomenon of hybrid vigour was noticed in it by Hedrick and Booth (1907). Though present day tomato is a self pollinated crop the unusual high heterosis observed in this has been attributed the fact that, originally tomato was a highly out crossing genus which was later evolved into a self pollinated one (Rick, 1965). Further, comparative ease in emasculation, high percentage of fruit setting and good number of seeds per fruit lead to the exploitation of heterosis in tomato. Though lot of information is available

on heterosis in tomato, it holds promise for further investigation, for future use whenever needed. It is the established fact that for successful hybrid breeding (development) selection of parents has vital role. The studies on combining ability of parents is essential in choosing parents. Many biometerical procedures have been developed to obtain information on combining ability and diallel crossing technique is one among them which is widely used to study combining ability of the parents to be chosen for heterosis breeding. Further, it is useful in providing information on gene action which is of great value to plant breeder in chosing the most advantageous breeding or selection procedure for improvement of attributes in question. Keeping aforesaid information in view a study on tomato was initiated with the following objectives. 1. To compare the variability for yield and other yield related traits in the populations generated by inter and intra population mating 2. To assess the variability for yield and other yield related traits in selfed and biparental populations 3. To study the shift in association pattern of component traits with yield among biparental populations 4. To study the magnitude of heterosis for growth, yield and quality parameters using diallel mating design and 5. To estimate combining ability effects and variances for yield and quality component characters

II. REVIEW OF LITERATURE

The main goal of most of the plant breeding programme is to increase the yielding ability of crop plants. Along with information on variability, the information on heterosis and combining ability of various quantitative and quality traits is also essential for crop improvement. This calls for taking stock of earlier works on crop improvement aspects of tomato. Since, there are no reports available on biparental mating work in tomato, the literature on variability, correlation and path analysis studies made in BIPs in crops like wheat and safflower is presented below. The literature on heterosis and combining ability studies in tomato is also presented below, under following headings. 2.1 Biparental mating 2.2 Diallel mating

2.1 BIPARENTAL MATING (BIP)

The review of literature concerning the studies conducted for this dissertation is outlined under the following headings. 2.1.1 Relative efficiency of BIPs over selfing series

2.1.2 Correlation co-efficient and path co-efficient analysis in segregating populations

2.1.1 Relative efficiency of biparental mating (BIPs) over selfing series

Tomato is predominantly a self pollinated crop. So presence of linkage blocks and inverse relations among the correlated characters are more common. Further the existing breeding procedures have the weakness of causing rapid homozygosity, low genetic variability, poor recombination rate and negative correlation among yield components (Clegg et al., 1972). Under such circumstances, the biparental mating in an appropriate segregating population is effective in breaking larger linkage blocks mostly in repulsion phase and provides more chances of recombination than selfed family selection (Gill, 1987). It is a useful system of mating for generation of increased variability and may appropriately be developed where lack of desired variation is the immediate bottleneck in the breeding programmes. It also avoids the early fixation of genes in homozygous state and provides a greater variability for selection to be effective for longer period. It has been possible to obtain additional variability for seed yield and other yield contributing characters from biparental mating in wheat and safflower (Yunus and Paroda, 1983 and Parameshwarappa et al., 1997) and other crops. In tomato, so far there have been no reports on the usefulness of biparental mating. Hence, the literature available on various aspects with respect to biparental mating and selfed generations in wheat and safflower is reviewed and presented under the following headings and explained characterwise. 2.1.1.1 Variability parameters in wheat A brief review of literature available on variability parameters in BIPs and selfed populations of wheat is summarised in Table 1 and explained here under. 2.1.1.1.1 Mean and Range Comparison of mean and range of different characters between biparental (BIPs) and selfed progenies indicated that in general, mean and range values of BIPs were higher than that of selfed progenies for all the characters studied (Nand et al., 1990a, Nematullah and Jha 1993 and Sharma et al. 1995). The mean and range pertaining to different characters in BIPs as well as selfed series are given in Table and are explained here under. Plant height Among the nine available reports on mean for plant height only four showed higher mean value in selfed series. Out of seven reports on range, two reports suggested wider range in BIPs while remaining indicated wider range in selfed populations. However, in most of the cases in BIPs, lower limit was slightly greater than selfed populations.

Table 1. Review of literature on comparison of variability, heritability and genetic advance for various characters in BIPs and selfed population in wheat

Character 1. Plant height (cm) Mean Selfed 109.24 Low 108.2 Low 82.5 114.4 114.5 108.8 111.33 2. Tillers per plant 9.06 Low Low 10.0 11.4 11.2 10.2 8.3 8.9 BIPs 110.96 High 102.8 High 77.0 105.9 104.2 97.7 129.86 9.72 High High 12.7 12.3 14.2 15.4 8.3 10.9 89.0 134.4 75.3128.9 Narrow 43-106 101.2124.5 90.2130.4 103.33113.33 7.2-13.2 Narrow 4.35 10.15 5.2-16.8 5.11 4.4-19.8 Range Selfed BIPs 89.8133.4 85.2112.7 Wider 43-104 98.2119.8 88.1108.5 100.0150.67 7.613.2 Wider 7.37 6.20 8.421.3 5.11 5.415.0 PCV Selfed 9.10 10.8 Low 12.6 5.2 7.24 18.83 17.8 Low 36.7 21.5 20.6 22.2 BIPs 7.50 9.5 High 12.4 11.8 11.7 13.53 15.8 High 42.5 18.2 15.6 13.8 GCV Selfed 7.65 9.8 12.7 Low 10.1 11.4 6.69 8.85 11.5 Low 25.6 15.9 14.8 17.6 BIPs 4.95 8.2 9.2 High 8.3 11.18 2.34 9.7 High 30.9 14.2 11.3 12.2 Heritability Selfed 71 81.9 79.4 Low 64.3 85.32 3 41.8 Low 50.0 55.2 63.3 BIPs 44 75.8 44.5 High 45.2 90.63 22 37.2 High 53.1 60.9 77.9 14.16 15 Low 37.6 21.4 2.6 18 25.3 Low 16.8 GA Selfed BIPs 15 13.1 High 11.5 28.46 12 High 46.5 22.9 2.4 Reference Randhawa and Gill (1978) Yunus and Paroda (1998) Gurudev Singh et al. (1986) Singh et al. (1988) Nanda et al. (1990a) Nanda et al. (1990b) Powar et al. (1990) Nematullah and Jha (1993) Sharma et al. (1995) Randhwa and Gil (1978) Yunus and Paroda (1983) Singh et al. (1988) Shrivastava et al. (1989) Nanda et al. (1990a) Nanda et al. (1990b) Nematullah and Jha (1993) Kaushik et al. (1996) Kaushik et al. (1996)

Table 1. Contd.....

Mean Selfed 3. Spike length 10.98 Low Low 11.9 12.8 4. Spikelet per spike 21.9 Low Low 20.1 63.3 5. Grains per spike 54.81 Low 60.3 BIPs 11.37 High High 13.2 13.3 22.18 High High 24.4 68.3 57.73 High 63.3 Range Selfed 9.3-13.8 Narrow 9.2-13.6 9.1-16.6 18.824.8 Narrow 17.824.6 50.072.3 37.367.5 54.369.7 BIPs 10-13.7 Wider 101.114.3 10.620.0 17.923.6 Wider 18.925.5 5.781.0 37.368.1 58.874.9 PCV Selfed 8.40 9.5 Low 8.8 6.23 6.7 Low 9.4 11.57 15.0 BIPs 6.52 8.6 High 8.3 5.02 6.7 High 10.2 9.85 13.7 GCV Selfed 5.86 6.4 Low 5.9 4.36 5.5 Low 7.2 6.86 10.7 7.1 BIPs 4.01 6.3 High 7.1 2.22 5.1 High 7.9 2.06 10.7 101.1 Heritability Selfed 49 44.3 Low 45.5 20 66.1 Low 58.3 35.0 50.4 36.2 BIPs 38 54.1 High 58.3 49 57.7 High 60.4 4.0 61.1 62.3 GA Selfed 9 Low 8.3 9 Low 11.3 16.0 5.4 BIPs 10 High 9.9 8 High 12.7 17.0 1.04 Randhawa and Gill (1978) Yunus and Paroda 91983) Singh et al. (1988) Nanda et al. (1990b) Nematullah and Jha (1993) Randhwa and Gill (1978) Yunu and Paroda (1983) Singh et al. (1988) Nanda et al. (1990b) Nematullah and Jha (1993) Randhawa and Gill (1978) Yunus and Paroda (1983) Gurudev Singh et al. (1986)

Character

Reference

Table 1. Contd.....

Character Mean Selfed Low 45.0 51.8 60.2 44.3 63.3 44.28 6. 1000-grain weight (g) 37.7 Low 39.2 Low 27.0 36.8 37.0 40.6 40.8 44.19 BIPs High 53.8 58.8 64.8 55.2 68.3 56.1 39.8 High 92.0 High 29.4 48.5 41.0 48.7 41.9 52.36 Range Selfed Narrow 20-70 43.67 52.369.6 50.072.3 22.767.17 41.772.5 36.241.8 Narrow 13.231.4 25.248.7 31.050.0 35.146.4 39.8357.67 BIPs Wider 26-69 38.83 54.972.1 57.081.0 26.1773.73 42.766.6 36.842.1 Wider 20.252.4 30.851.9 34.051.0 37.747.4 36.9365.17 Low 6.9 7.2 19.7 26.79 10.04 11.4 Low 10.5 3.24 15.7 10.91 PCV Selfed BIPs High 9.9 7.5 13.2 23.79 9.08 9.3 High 10.3 32.7 11.3 12.57 Low 5.7 5.7 12.4 26.24 7.92 6.9 1.4 Low 8.7 24.3 11.8 9.45 GCV Selfed BIPs High 7.2 5.7 8.2 22.08 7.29 7.1 2.3 High 9.0 26.5 7.9 11.52 Heritability Selfed Low 75.7 56.2 95.9 62.0 34.7 23.1 Low 68.9 56.3 75.03 BIPs High 51.9 61.5 86.10 65.0 57.7 33.3 High 76.5 61.1 84.02 Low 10.3 8.9 23.44 8.0 0.5 Low 14.9 37.6 7.45 GA Selfed BIPs High 10.6 9.2 18.81 11.0 1.1 High 16.3 41.2 11.39 Randhaw and Gill (1978) Yunus and Paroda (1983) Gurudev Singh (1986) Singh et al. (1988) Singh et al. (1988) Shrivastava et al. (1989) Nanda et al. (1990b) Pawar et al. (1990) Nematullah and Jha (1993) Sharma et al. (1995) Reference Singh et al. (1988) Srivatava et al. (1989) Nanda et al. (1990a) Nanda et al. (1990b) Pawar et al. (1990) Jha (1993)

Table 1. Contd.....

Character Mean Selfed 41.6 7. Grain yield per plant (g) 14.69 Low 23.8 Low 26.3 13.4 20.3 117.87 11.6 BIPs 43.6 16.77 High 27.3 High 33.6 16.1 26.5 106.62 20.2 Range Selfed 29.149.2 14.128.4 11.732.9 Narrow 10.560.3 7.0-21.0 67.0161.7 BIPs 37.850.3 15.732.5 12.336.7 Wider 12.072.3 6.230.3 65.0175 6.3 24.6 21.2 Low 27.6 41.1 16.49 28.4 PCV Selfed BIPs 5.8 14.98 19.8 High 28.3 36.8 29.10 17.9 5.1 5.98 11.4 14.1 Low 17.6 28.9 16.19 24.1 GCV Selfed BIPs 5.3 9.54 13.3 16.7 High 18.2 26.2 28.59 16.4 Heritability Selfed 65.9 6.0 29.1 31.7 Low 40.2 96.42 72.2 BIPs 84.0 42.0 45.1 48.9 High 51.6 96.52 83.9 3.6 13.0 4.3 Low 22.9 38.6 4.9 GA Selfed BIPs 4.4 18.0 6.6 High 27.1 61.71 6.3 Reference Kaushik et al. (1996) Randhawa and Gill (1978) Yunus and Paroda (1983) Gurudev Singh et al. (1986) Singh et al. (1998) Srivastava et al. (1989) Nanda et al. (1990a) Powar et al. (1990) Sharma et al. (1995) Kaushik et al. (1996)

Tillers per plant Out of nine reports on tillers per plant, higher mean value was noticed in all the reports for this trait except in one where mean value was equal in BIPs as well as selfed population. Of the available seven reports on range only three showed wider range in BIPs, three indicated wider range in selfed populations while the remaining one suggested equal range in both the methods. Spike length All the five reports available on this trait showed higher mean value in BIPs when compared to selfed series. Out of four reports available on range, two reports suggested wider range in BIPs and the other two indicated wider range in selfing series. But the lower limit in BIPs in all the reports was higher than the corresponding value in selfed progenies. Spikelet per spike There are five reports available on mean for this trait, in all the cases, increase in mean value has been observed in BIPs over their selfed progenies. With regard to range two reports showed wider range in BIPs and though range was narrow in BIPs in the other two reports, the lower and upper limits were higher in BIPs when compared to corresponding selfed population. Grains per spike All the eleven reports indicated higher mean value for this trait in BIPs when compared to corresponding selfed population. Out of nine reports, seven reports indicated wider range in BIPs while only two suggested wider range in selfed population. 1000-grain weight It is worth while to note that, all the 11 reports available on mean 1000 grain weight showed higher mean in BIPs over their selfed generations. But this trend was not seen in case of range for this trait. Of the nine reports only three reports wider range was observed in BIPs while in the remaining eight reports, wider range was seen in selfed progenies. Grain yield per plant Out of eleven reports except one all the other ten reports showed higher mean values in BIPs when compared to corresponding selfed progenies. 2.1.1.1.2 Variance Although, mean and range values provide preliminary idea about the variability, coefficient of variation is a reliable unit of measurement. The review indicated that the character having higher range normally exhibited higher genotypic and phenotypic co-efficient of variation. The coefficient of variations were slightly higher in BIPs when compared to the selfed progenies for most of the characters studied (Srivastava et al., 1989). Plant height Out of seven reports on variability (PCV and GCV), three each suggested low and moderate variability for this trait in BIPs. The same trend was also reported in selfed progenies for this character. However, only one report revealed higher variability in BIPs whereas in corresponding selfed population, it was low. Tillers per plant There are seven reports available on variability of this trait. Out of them, five reports indicated moderate variability in BIPs and the remaining two reports suggested high variability. On the other hand, one, two and four reports revealed low, moderate and high variability, respectively in the corresponding selfed progenies. Spike length Among the four reports available on variability with respect to spike length, only one report indicated high variability and the remaining three suggested low variability in BIPs whereas, in corresponding selfed series, all the reports suggested low variability for this character.

Spikelets per spike Of the four reports, two reports suggested low variability while one report each indicated moderate and high variability, respectively for this trait in BIPs. On the contrary, all the reports showed low variability in selfed population. Grains per spike Of the available nine reports, four indicated low, three suggested moderate and the remaining two noticed high variability in BIPs whereas in selfing series five reports suggested low, three reports revealed moderate and the remaining one showed high variability for this trait. 1000-grain weight Out of nine reports on variability for this trait, four indicated presence of high variability in BIPs. On the other hand, three suggested low, five indicated moderate and the remaining one suggested high variability in the corresponding selfed progenies. Grain yield per plant Among the nine available reports on variability, four indicated moderate and the remaining five suggest high variability in BIPs. Where as one, two and six indicate low, moderate and high variability for this trait in the corresponding selfed series. 2.1.1.1.3 Heritability and genetic advance The co-efficient of variation indicate only the extent of total variability present for a character and do not demarcate it into heritable and non-heritable portion, where as the heritability computed in broad sense would suggest how far the variation is heritable and consequently selection is effective. High estimate of heritability coupled with high genetic advance is more helpful in accurately predicting genetic gain under selection than heritability estimate alone (Johanson et al., 1955). A review of the studies pertaining to heritability (%) and genetic advance in BIPs and selfed population is presented in Table 1 and summarized here below. Plant height Among the available six reports on heritability, three each suggested moderate and high heritability respectively in BIPs while in selfed progenies, high estimate of heritability was observed in all the reports except one where low heritability was observed. With regard to genetic advance, out of the five reports, three indicated moderate while two suggested high heritability estimate in BIPs. In selfed series, except one report all suggested moderate genetic advance. Tillers per plant Out of six reports on heritability one showed low, two indicated moderate and the remaining three suggested high heritability estimate for this trait in BIPs, whereas in selfed populations, heritability estimate was low, moderate and high as indicated by two, three and one reports respectively. With regard to genetic advance, of the five available reports, one report each indicated low and moderate estimates and the remaining three suggested high genetic advance for this trait in BIPs. On the contrary, two reports indicated low, one suggested moderate and other two observed high genetic advance in the corresponding selfing series. Spike length In BIPs, except one, which indicated high heritability, the remaining three suggested moderate heritability. In selfed populations three reports indicated moderate while one suggested low heritability estimate. With respect to genetic advance, only three reports are available, in which one each indicated low, moderate and high genetic advance in BIPs, while in selfed populations, low heritability estimates were recorded. Spikelets per spike Among four reports, two each indicated moderate and high heritability estimate in BIPs whereas in selfed progenies two suggested low, one moderate and the remaining one

suggested high heritability estimate for this trait. Out of three reports on genetic advance, one report each indicated low, moderate and high estimate in BIPs. However two suggested low and one report indicated moderate genetic advance in selfed populations. Grains per spike Among eight available reports, only one report each indicated low and moderate heritability estimates while the remaining six reports revealed high estimate for this trait in BIPs. On the other hand, one, four and three reports indicated low, moderate and high heritability estimate respectively in the corresponding selfed progenies. Out of seven reports on the genetic advance, two suggested low, four moderate and the remaining one indicated high estimate in BIPs whereas in selfed generations, four suggested low, two moderate and the remaining one suggested high estimate of genetic advance. 1000-grain weight There are eight reports on heritability estimate for this trait, of which except two which indicated moderate, the remaining six reports suggested high heritability estimate. With respect to genetic advance, out of seven reports, two indicated low and three showed moderate estimate of genetic advance, while the remaining two suggested high genetic advance in BIPs. In selfed progenies five reports indicated low while one report each suggested moderate and high genetic advance estimate for this trait. Grain yield per plant Of the available eight reports, four reports indicated moderate heritability estimate while the remaining four reports suggested high estimate of heritability in BIPs. In selfed generations, three reports suggested presence of low, two indicate moderate and three suggested high heritability estimates. With regard to genetic advance out of seven reports two suggested low, one as moderate and the remaining four reports indicated high genetic advance in BIPs for this trait. In selfed population, three reports indicated low, one moderate and the remaining three suggested high genetic advance for this trait. 2.1.1.2 Variability parameters in safflower A brief review with regard to variability in BIPs and selfed generations in safflower is given in Table 2 and is presented here under. 2.1.1.2.1 Mean and range Plant height All the three available reports suggested high mean values in selfed progenies as compared to BIPs. As regard to genetic variability, of the three reports, two indicate wider range in BIPs and one suggested narrow range when compared to selfed generations. Number of branches Of the four reports available on mean and range for number of branches, all the four suggested higher mean. These three indicated wider range while one report recorded narrow range in BIPs as compared to selfed populations. Number of capitula Of the four reports three indicated higher mean and one lower mean. Two reports suggested wider range while the other two indicated narrow range for this character in BIPs as compared to selfed populations. Size of capitula Out of four reports, three suggested higher mean value and the other one suggested low mean value. All the four reports indicated wider range for this trait in BIPs when compared to selfed progenies. Number of seeds per capitula One report showed lower mean, while three reports indicated higher mean values in BIPs. Three reports suggested wider range in BIPs and one indicated equal range both in selfs and Bips.

Table 2. Review of literature on comparison of variability, heritability and genetic advance for various character in BIPs and selfed populations in safflower

Character 1. Plant height (cm) Mean Selfed 112.30 121.50 74.57 2. Number of branches 6.40 11.20 9.00 9.80 3. Number of capitula 16.50 15.50 18.48 21.28 BIPs 110.70 105.60 71.81 7.50 13.50 11.06 10.55 13.50 17.20 21.84 26.64 Range Selfed 114.7131.0 101.5141.6 66.0083.50 5.408.80 7.2013.40 6.7512.20 7.0012.00 7.6024.90 6.6034.10 14.1027.95 16.0024.00 BIPs 91.5132.5 95.6145.62 64.0080.00 5.509.70 7.3014.90 8.512.70 8.0013.50 11.9038.10 7.8031.70 15.6025.80 19.0032.00 PCV Selfed 8.39 16.37 17.20 BIPs 7.68 16.04 19.95 GCV Selfed 0.67 3.17 6.65 10.01 8.14 11.63 20.85 18.33 14.46 BIPs 6.40 22.4 6.97 14.36 19.02 14.90 28.47 33.57 13.19 Heritability Selfed 2.51 45.84 62.80 56.16 75.61 50.50 50.43 80.81 41.80 BIPs 70.79 92.91 82.40 69.50 86.67 86.40 68.69 90.48 54.00 GA Selfed 0.25 5.36 10.86 0.98 1.63 17.00 5.03 5.26 16.58 BIPs 12.29 47.06 13.02 1.85 4.92 28.53 6.65 11.31 19.96 Reference Singh and Sahu (1981) Singh and Sahu (1981) Kadlera (1997) Singh and Sahu (1981) Singh and Sahu (1981) Parmeshwarappa et al. (1997) Kadlea (1997) Singh and Sahu (1987) Singh and Sahu (1987) Parameshwarappa et al. (1997) Kadlera (1997)

Table 2. Contd....

Mean Selfed 4. Size of capitula 2.60 2.10 2.18 2.28 5. Number of seeds per capitula 41.50 26.60 30.78 35.11 6. 100 seed weight (g) 5.00 4.00 5.11 5.07 BIPs 2.90 2.40 2.16 2.34 59.30 32.70 33.40 34.43 5.50 5.00 4.41 5.24 Range Selfed 2.303.00 2.002.30 2.052.35 2.072.40 13.7058.40 8.5041.50 24.0036.50 24.0047.00 4.906.20 2.505.80 4.25 4.096.75 BIPs 2.203.90 2.002.50 1.852.38 2.152.51 18.2101.6 8.2051.00 27.0041.00 24.0047.00 4.908.10 2.506.30 4.05 3.756.60 PCV Selfed 5.60 21.84 17.54 BIPs 5.81 23.81 16.32 GCV Selfed 2.11 3.01 4.90 18.83 21.80 17.99 14.21 16.82 BIPs 7.56 11.93 5.30 16.63 27.89 22.33 5.55 15.55 Heritability Selfed 42.86 50.00 76.60 93.58 73.57 67.80 7.01 92.00 BIPs 81.36 94.53 83.30 70.25 83.04 87.90 75.23 90.80 GA Selfed 0.10 0.12 8.7 15.57 10.25 30.50 0.54 33.33 BIPs 0.41 0.57 9.82 17.08 17.12 43.30 0.38 30.53 Singh and Sahu (1981) Singh and Sahu (1981) Parameshwarappa et al. (1997) Kadlera (1997) Singh and Sahu (1981) Singh and Sahu (1981) Parameshwarappa et al. (1997) Kadlera (1997) Singh and Sahu (1981) Singh and Sahu (1981) Paramesharappa et al. (1997) Kadlera (1997)

Character

Reference

Table 2. Contd....

Mean Selfed 7. Grain yield per plant (g) 19.30 12.40 52.20 23.02 8. Oil content (%) 30.10 32.30 31.40 28.61 BIPs 21.50 19.50 26.18 31.98 32.70 35.70 33.32 29.69 Range Selfed 9.4024.80 8.401860 16.5032.00 17.0032.20 26.6033.80 27.4035.20 28.9032.70 26.8930.28 BIPs 14.1033.70 9.5021.40 21.9031.00 23.0039.00 28.7036.40 29.8039.20 31.5034.60 26.2032.30 PCV Selfed 3.35 4.16 BIPs 5.36 5.05 GCV Selfed 13.71 15.98 3.34 4.24 3.62 BIPs 21.58 22.08 5.28 6.20 4.81 Heritability Selfed 74.79 58.84 99.60 61.99 75.80 BIPs 79.37 82.08 97.10 70.62 90.90 GA Selfed 4.70 3.13 30.17 2.07 6.88 BIPs 8.52 8.04 32.98 3.51 10.71 Singh and Sahu (1981) Singh and Sahu (1981) Parameshwarappa et al. (1997) Kadlera (1997) Singh and Sahu (1981) Singh and Sahu (1981) Parameshwarappa et al. (1997) Kadlera (1997)

Character

Reference

100-seed weight Out of four available reports on mean and range, three suggested high while one indicated low mean. All the four suggested wider range for this character in BIPs over their selfs. 2.1.1.2.2 Variability, heritability and genetic advance High variability, heritability and genetic advance have been reported for most of the characters in BIPs when compared to corresponding selfed generation (Singh and Sahu, 1981 and Kadlera, 1997). A review of earlier reports on these aspects is summarized in Table 2 and explained character wise here under. Plant height Out of three reports on variability for this character, one indicated high and two suggested low variability in BIPs whereas in selfed progenies all the three reports suggested low variability. With respect to heritability, all the available three reports indicates high estimate in BIPs while one each for low, moderate and high heritability is available for selfed progenies. Of the three reports on genetic advance, two indicated moderate estimate while one indicated high genetic advance in BIPs whereas except one which is moderate, remaining two suggested low estimate in selfed series. Number of branches Among the three reports available on variability, heritability and genetic advance, all suggested moderate variability for this character in BIPs while two reports suggested moderate and one indicated low variability in corresponding selfed populations. With respect to heritability, high estimate was reported by all the three workers in BIPs whereas only one suggested high estimate and the remaining two indicated moderate heritability in selfed generations. With regard to genetic advance one report indicated high estimate and these indicated low estimates in BIPs. In selfed populations low and moderate genetic advance estimates were reported. Number of capitula Among the available three reports on variability and genetic advance, two indicated high and the remaining one suggested moderate variability in BIPs while in selfs, two reports suggest moderate and one high estimate. With respect to heritability in BIPs, two reports indicated high while the remaining one suggested moderate heritability. In selfed progenies, moderate and high heritability was reported. With respect to genetic advance, two indicated moderate while the other indicated low estimate in BIPs whereas, two and one report suggested low and moderate estimate respectively in selfed populations. Size of capitula There are three reports available on variability, heritability and genetic advance. Out of these in BIPs, one suggested moderate variability and two reports indicated low variability. In the corresponding selfed progenies, low variability was observed in all the reports for this character. With respect to heritability, all the reports suggested high heritability in BIP populations whereas in selfed populations, two reports suggested moderate and one indicated high heritability. With respect to genetic advance, low genetic advance estimate was reported in BIPs as well as in their selfed progenies. Number of seeds per capitula Of the three reports available on variability, heritability and genetic advance, two and one report suggested high and moderate variability, respectively in both BIPs as well as selfed series. With regard to heritability, both populations showed high heritability for this character. On the other hand one suggested high and the remaining two indicated moderate genetic advance estimate in both the populations.

Table 3. Review of literature on correlation studies in BIPs and selfed population in wheat Character 1. Plant height (cm) Positive Positive and significant Positive and significant Positive and significant Negative and significant Positive and significant 2. Tillers per plant Positive and significant Positive and significant Positive and significant Positive and significant Positive and significant Positive and significant Positive and significant 3. Spike length (cm) Positive and significant Positive and significant Positive Positive and significant Positive and significant Correlation with seed yield BIPs Positive Positive Positive and significant Positive and significant Negative Positive and significant Positive and significant Positive and significant Positive and significant Positive and significant Positive and significant Positive and significant Positive Positive and significant Positive and significant Positive and significant Positive Positive and significant Selfed Reference Randhawa and Gill (1978) Yunus and Paroda (1982) Singh and Misra (1984) Nanda et al. (1990a) Nanda et al. (1990b) Nematullah and Jha (1993) Randhawa and Gill (1978) Yunus and Paroda (1982) Singh and Misra (1984) Nanda et al. (1990a) Nanda et al. (1990b) Nematullah and Jha (1993) Subhash Chander et al. (1995) Randhawa and Gill (1978) Yunus and paroda (1982) Singh and Misra (1984) Nanda et al. (1990) Nematullah and Jha (1993)

Table 3. Contd.....

Character 4. Spikelets per spike

Correlation with seed yield BIPs Positive and significant Positive and significant Positive and significant Positive and significant Selfed Positive and significant Positive and significant Positive and significant Positive and significant Positive and significant Positive and significant Positive and significant Positive and significant Positive Positive and significant Positive and significant Positive and significant Positive and significant Positive and significant Positive Positive and significant Positive and significant Positive and significant

Reference Randhawa and Gill (1978) Yunus and Paroda (1982) Nanda et al. (1990a) Nematullah and Jha (1993) Randhawa and Gill (1978) Yunus and Paroda (1982) Singh and Misra (1984) Srivastava et al. (1989) Nanda et al. (1990a) Nanda et al. (1990b) Nematullah and Jha (1993) Randhawa and Gill (1978) Yunus and Paroda (1982) Singh and Mishra (1984) Srivastava et al. (1989) Nanda et al. (1990a) Nanda et al. (1990b) Nematullah and Jha (1993)

5. Grains per spike

Positive and significant Positive and significant Positive and significant Positive and significant Positive Positive and significant Positive and significant

6. 1000-grain weight (g)

Positive Positive Positive and significant Positive and significant Positive and significant Positive and significant Positive and significant

Grain yield per plant Of the three reports available on variability, heritability and genetic advance one suggested low and the other two indicated high variability in BIPs. But two reports suggested moderate and one indicated low estimate in selfed populations. With regard to heritability, all the three reports suggested high heritability in BIP populations whereas in selfed populations, one and two reports respectively indicated high and moderate heritability. On the other hand, two and one report indicated low and high genetic advance estimate, respectively in both the populations. Oil content Both the available reports on variability, heritability and genetic advance, suggested low variability in both selfed and intermated populations. Similarly the same two reports suggested high heritability in both populations. With respect to genetic advance, one indicated low and the other moderate genetic advance estimate in BIP population whereas both the reports suggested low genetic advance estimate for this character in selfed populations. 2.1.1.3 Correlation co-efficient analysis in wheat Correlation co-efficient analysis indicates the mutual relationship among various characters. The release of concealed variability as a result of biparental mating can also be examined by studying changes in direction and magnitude of correlations among different characters in BIPs and selfed progenies. The changes that occur in the correlations between BIPs and selfed progenies in various studies is given in Table 3 and are explained character wise here under. Plant height There are about six reports on correlation of plant height with seed yield. Out of six, four suggested positive and significant association, one indicated just positive correlation with seed yield in BIPs. In the corresponding selfed progenies both positive and negative association was exhibited. Tillers per plant All the seven reports indicated positive significant association of tillers per plant with seed yield in BIPs. In selfed populations, except one each for positive and negative significant association, remaining all indicated significant positive association between these two traits. Spike length Out of five reports, one indicated just positive while the remaining four reports indicated significant positive association of spike length with seed yield in BIP population. The same trend was also observed in selfed progenies. Spikelets per spike All the four available reports suggested positive and significant association of spikelets per spike with seed yield in BIPs as well as in selfed populations. Grains per spike Among the eight reports on correlation of grains per spike with seed yield, all the studies indicated positive and significant association except in two reports where positive association has been observed in BIPs. In selfed series, three reports indicated positive correlation, while the remaining five reports suggested positive and significant association with seed yield. 1000-seed weight in grams Out of seven reports available two indicated simple positive association while the remaining five suggested positive and significant association with seed yield in BIPs. In selfed progenies, except one report suggesting positive association, the other suggested positive significant association with seed yield.

2.1.1.4 Correlation studies in safflower Plant height (cm) appeared to be positively and significantly associated with seed yield in selfed population while it was non-significant in BIPs (Kadlera, 1997). Number of branches has been reported to be associated positively and significantly with seed yield (Kadlera, 1997) in selfed as well as in BIPs. However, in BIPs its magnitude of association with seed yield changed to non-significant (Parameshwarappa et al., 1997). It has been observed that number of capitula per plant had positive and significant correlation with seed yield in BIPs (Parameshwarappa et al., 1997 and Kadlera 1997) while in selfed population, its association was changed to negative (Kadlera, 1997). The positive association of capitula size (cm) with seed yield in BIPs was changed in its direction in selfed progenies (Parameshwarappa et al., 1997). Kadlera (1997) reported negative significant association in selfed progenies whereas in BIPs, its magnitude only reduced to become non-significant. Number of seeds per capitula showed positive correlation with seed yield in BIPs and selfed progenies by Parameshwarappa et al. (1997), however negative association in selfs was reported by Kadlera (1997). Parameshwarappa et al. (1997), observed positive and strong association of 100 seed weight with seed yield in BIPs and weak positive association in selfed population, whereas, vice-versa was reported by Kadlera (1997). The significant negative association between oil content (%) and seed yield in selfs was reduced in its magnitude but not in direction and became non-significant with seed yield in BIPs (Parameshwarappa et al., 1997 and Kadlera, 1997). 2.1.1.5 Path co-efficient analysis in wheat and safflower Literature with regard to path coefficient analysis in BIPs is very meagre . A comparison of direct and indirect effects of various characters on seed yield in selfs and the BIPs, revealed that the changes in the nature and degree of association amongst various characters were accompanied by the change in their direct and indirect effects (Yunus and Paroda, 1982). In selfed progenies, plant height, tillers per plant, spike length (cm) had higher positive direct effect on seed yield when compared to BIPs (Yunus and Paroda 1982 and Nematullah and Jha, 1993). Yunus and Paroda (1982) reported high negative direct effect of spikelets per spike on seed yield in selfed generation as compared to BIP population. Grains per spike exhibited high positive direct effect in selfed population when compared with BIPs (Yunus and Paroda, 1982). Negative direct effect was observed by Nembatullah and Jha (1993) in selfed progenies as compared to the positive direct effect in BIPs. The 1000-grain weight had positive direct effect on seed yield in selfed generations and it changed to negative side in BIPs (Yunus and Paroda, 1982). 1000 grain weight showed highest positive direct effect on seed yield in BIPs when compared to selfs as reported by Nematullah and Jha, 1993). According to Yunus and Paroda (1982), highest positive direct effect of tillers per plant on seed yield was observed when compared to other characters. Positive direct effect of 1000 grain weight was high in BIP's than in selfed population when compared to other characters. Therefore by selecting for bolder grains in BIPs the grain yield can be improved (Nematullah and Jha, 1993). In safflower, only one available report on path analysis indicated number of seeds per capitula to have highest positive direct effect followed by 100 seed weight in selfs whereas, capitula weight (g) followed by number of branches recorded highest positive direct effect on seed yield (Kadlera, 1997) in BIPs.

2.1.2 Correlation co-efficient and path co-efficient analysis in segregating populations of tomato Association of economically important quantitative characters which is statistically determined by correlation coefficient has been quite useful as a basis of selection. Correlation studies provide information that the selection for one character will result in progress for all the correlated characters. Thus the review pertaining to the correlation among the different traits is presented here under. 2.1.2.1 Association of yield with other characters Positive correlation between yield and number of primary branches per plant was observed by Supe and Kale (1982), Manivannan and Irulappan (1986), Reddy and Lal (1990) Anandgouda (1997) and Patil (1998). The number of fruiting clusters per plant showed a positive relationship with yield in F1 and F2 generations as indicated by Kanthaswamy et al. (1994). Ponnuswamy and Muthukrishnan (1977) observed positive correlation between number of fruits per plant and yield. However, high positive correlation between yield per plant and number of fruits per plant was reported by Patil (1998). Dudi and Kalloo (1982) reported higher gentoypic correlation coefficient than the phenotypic correlations. Total yield exhibited positive correlation with number of fruits per plant. Similar results were reported by Nandpuri et al. (1973) and Prasad and Prasad (1977). Positive association was noticed between number of fruits per plant and yield by several workers (Nandpuri et al. 1973 and 1976; Prasad and Prasad, 1977; Bangaru 1981; and Patil 1998). A positive correlation between pericarp thickeness and yield was observed by Bhutani and Kalloo (1989) and Anandgowda (1997), Patil (1998). Kanthaswamy et al. (1994) reported positive correlation between TSS and yield, whereas, Anandgowda (1997) reported negative association for the same characters. Bhutani and Kallo (1989) observed negative correlation between yield and acidity, while Supe and Kale (1992), Rathod (1997) and Patil (1998) observed positive association. 2.1.2.2 Association among the characters A positive correlation between number of primary branches per plant and number of fruits per plant was observed by Supe and Kale (1992) and Rathod (1997). Rathod (1997) and Indunair and Thamburaj (1995) reported negative association of number of fruits per plant and size. 2.1.2.3 Path analysis The study of simple correlation do not provide an exact picture of relative importance of direct and indirect influence of each of the component character towards the direct character. So, this can be overcome by following path coefficient analysis technique by further partitioning the correlation coefficient into direct and indirect effects. Supe and Kale (1992) and Sonone et al. (1987) reported that the number of primary branches per plant had highest positive direct effect as the major yield contributing trait for enhancing yield. But, Patil (1998) noticed indirect effect of number of branches per plant through number of fruits per plant on yield was high. Number of fruits per plant had highest positive direct effect on yield (Padda et al., 1971, Bhutani and Kalloo, 1989, Rathod, 1997 and Patil 1998). Bhutani and Kalloo (1989) reported that number of locules per fruit had direct positive effect on yield but its effect was counter balanced by indirect negative effects via number of fruits. A negative and low indirect effects of pericarp thickness, TSS and per cent juice recovery on yield of fruits per plant was noticed through number of locules, as reported by Patil (1998). Pericarp thickness had positive direct effect on yield mainly due to positive indirect effects through number of fruits per plant and number of branches per plant (Patil, 1998).

Table 4. Heterosis for different traits in tomato as reported by various authors Characters Growth parameters 1. Plant height No. of crosses Range of heterosis (%) over Better parent Commercial check 53.73-36.33 1.90-41.10 0.42-45.37 -31.45-28.05 12.10-26.40 Positive -15.42-9.43 4.5-12.80 -5.27-5.63 7.40-40.10 4.65-13.06 -25.53-14.28 -50.98-60.45 12.74-13.94 -123.4-0.53 14.29-27.73 -23.29-22.23 -22.27-23.92 -58.82-148.64 -3.10-15.30 52.81-70.79 References

Mid parent 2.86-83.19 -14.73-74.11 Positive 6.5-16.56 -27.77-8.87 37.5-5.1 -8.37-24.05 -65.52-81.27 39.70-42.99 -6.22-6.51 -

28 F1 hybrids 6x6 diallel 8x8 diallel 30 hybrids 10 F1 hybrids 16 F1 hybrids 12 F1 hybrids 15 F1 hybrids 15 F1 hybrids 5 F1 hybrids 5 F1 hybrids 28 F1 hybrids 12x12 diallel 28 F1 hybrids 12x12 diallel 6 F1 hybrids 50 F1 hybrids 36 F1 hybrids 8x8 half diallel 91 F1 hybrids 45 F1 hybrids

Bhutani et al. (1973) Singh et al. (1976) Mishra and Khanna (1977) Yogananda (1977) Singh et al. (1978) Peter and Rai (1978) Anbu et al. (1981) Ahmed et al. (1988) Kumar et al. (1998) Ramamohan (1988) Kanthaswamy and Balakrishnan (1989) Dundi (1991) Dod et al. (1992) Singh and Singh (1993) Dod et al. (1995) Nagaraja (1955) Dharmatti (1995) Ghosh et al. (1997) Patil (1997) Bhatt et al. (1999) Fageria (2001)

Table 4. Contd..... Characters No. of crosses 9 F1 hybrids 8x8 diallel 28 F1 hybrids 8x8 diallel 16 F1 hybrids 10x10 diallel 15 F1 hybrids 5 F1 hybrids 5 F1 hybrids 28 F1 hybrids 12x12 diallel 12x12 diallel 28 F1 hybrids 50 F1 hybrids 36 F1 hybrids 8x8 half diallel 45 F1 hybrids 91 F1 hybrids 9 F1 hybrids Range of heterosis (%) over Better parent Commercial check -18.9-38.9 36.2-59.6 -34.54-53.30 -58.47-60.10 1.77-67.20 55.84-93.49 Positive 55.84-93.49 Positive -8.77-31.58 -6.70-15.67 16.006-57.237 -32.77-71.11 2.29-7.46 -3.22-6.47 -79.88-62.16 -76.07-231.29 22.54-28.74 -32.26-90.65 -84.39-4.28 -18.9-38.9 36.2-59.6 References Tiwari and Lal (2004) Thakur (2004) Bhutani et al. (1973) Mishra and Khanna (1977) Peter and Rai (1978) Sidhu et al. (1981) Patil (1984) Kumar et al. (1988) Ramamohan (1988) Kanthaswamy and Balakrishnan (1989) Dundi (1991) Dod et al. (1992) Dod et al. (1995) Singh and Singh (1993) Dharmatti (1995) Ghosh et al. (1997) Patil (1997) Shrivastava et al. (1998) Bhatt et al. (1999) Tiwari and Lal (2004)

2. Number of primary branches

Mid parent 25.60-50.80 5.71-76.00 Negative Positive 24.80-72.50 17.824-60.367 -12.66-12.26 -78.84-83.23 43.26-45.67 -2.99-46.02 66.57-122.16 25.6-50.8

B. Yield and yield components 1. Number of flowers per truss

45 F1 hybrids 16 F1 hybrids 8x8 half diallel

-25.03-17.23 53.45-19.15

-25.75-16.36 1.64-7.64 -

31.90-39.75 12.63-481.89

Tendulkar (1994) Peter and Rai (1978) Patil (1997)

Table 4. Contd..... Characters 2. Number of fruits per truss No. of crosses 16 F1 hybrids 28 F1 hybrids 28 F1 hybrids 45 F1 hybrids 10 F1 hybrids 50 F1 hybrids 8x8 half diallel 91 F1 hybrids 8x8 diallel 8x8 diallel 28 F1 hybrids 66 F1 hybrids 6x6 diallel 30 F1 hybrids 10 F1 hybrids (10x3) (LxT) analysis 15 F1 hybrids 15 F1 hybrids 5 F1 hybrids 105 F1s + 15 parents 28 F1 hybrids Range of heterosis (%) over Better parent Commercial check 57.78-66.92 87.06-96.55 -25.00-78.07 -54.59-17.80 -31.76-17.23 -49.39-11.87 10.07-21.07 -100.0-7.94 -100.00-278.35 -21.43-50.00 -65.68-0.36 -17.06-46.62 -46.83-49.53 -47.07-95.13 -37.88-313.63 2.00-54.40 -28.26-53.71 4.00-55.70 -45.82-81.28 Positive -31.54-27.47 -25.69-13.25 -23.31-48.67 9.33-57.24 -22.33-207.08 References Peter and Rai (1978) Dundi (1991) Singh and Singh (1993) Tendulkar (1994) Pujari and Kale (1994) Dharmatti (1995) Patil (1997) Bhatt et al. (1999) Thakur et al. (2004) Chaudhary and Khanna (1972) Bhutani et al. (1973) Rattan and Saini (1976) Singh et al. (1976) Yogananda (1977) Singh et al. (1978) Jamwal et al. (1984) Ahmed et al. (1988) Kumar et al. (1988) Kanthaswamy and Balakrishnan (1989) Yadav et al. (1989) Dundi (1991)

3. Number of fruits per plant

Mid parent -20.87-80.95 -8.70-16.88 -25.50-25.60 -100.0-40.62 -59.42-23.63 -6.25-81.36 -13.16-156.46 20.65-98.93 -

Table 4. Contd..... Characters No. of crosses 12x12 F1 hybrids 28 F1 hybrids 28 F1 hybrids 9 two parents + 42 thread parents + 18 four parents 45 F1 hybrids 12x12 diallel 50 F1 hybrids 6 F1 hybrids 36 F1 hybrids 8x8 half diallel 43 F1 hybrids 91 F1 hybrid 9 F1 hybrids 8x8 diallel 8x8 diallel 30 F1 hybrids 12 F1 hybrids 45 F1 hybrids 15 F1 hybrids 5 F1 hybrids Range of heterosis (%) over Better parent Commercial check -47.61-58.00 -30.3-114.88 References Dod et al. (1992) Singh and Singh (1993) Pujari and Kale (1994) Sundaram et al. (1994)

Mid parent -7.58-24.44 -9.24-134.06

4. Average fruit weight (g)

-17.38-114.19 -100-142.62 -1.03-51.73 47.15-55.96 31-23-37.79 58.16-74.15 5.90-248.9 -47.63-24.78 -62.86-15.5 -35.40-65.30 31.50

-34.10-96.89 10.78-46.51 -100-94.74 -40.51-0.83 30.94-53.51 -132.98-1.74 25.9-153.8 -17.41-29.89 -57.50-6.82 -79.19-9.68 67.28-88.39 Positive 31.50

-47.87-113.98 -100.00-1133.50 -40.59-33.75 15.9-137.5 -

Tendulkar (1994) Dod et al. (1995) Dharmatti (1995) Nagaraja (1995) Ghosh et al. (1997) Patil (1997) Shrivastava et al. (1998) Bhatt et al. (1999) Tiwari and Lal (2004) Thakur (2004) Chaudhary and Khanna (1972) Yoganand (1977) Anbu et al. (1981) Rajput (1987) Ahmed et al. (1988) Rammohan (1988)

Table 4. Contd..... Characters No. of crosses 105 F1 hybrids +15 parents 28 F1 hybrids 12x12 diallel 28 F1 hybrids 28 F1 hybrids 13x13 diallel 45 F1 hybrids 9 two parent + 42 three parent + 18 four parent 6 F1 hybrids 36 F1 hybrids 8x8 half diallel 45 F1 hybrids 9 F1 hybrids 8x8 diallel 28 F1 hybrids 66 F1 hybrids 6x6 diallel 30 F1 hybrids 16 F1 hybrids 10 F1 hybrids 12 F1 hybrids Range of heterosis (%) over Better parent Commercial check -29.11-25.70 -49.62-38.53 8.08-24.91 -30.47-76.45 -7.18-29.28 -51.13-8.71 -48.27-53.6 -72.40-114.95 References Yadav et al. (1989) Dundi (1991) Dod et al. (1992) Singh and Singh (1993) Pujari and Kale (1994) Reddy and Reddy (1994) Tendulkar (1994) Sundarm et al. (1994)

Mid parent -37.63-55.40 -6.16-31.54 -8.07-38.98 -36.21-22.95 -35.6-20.69

-52.58-62.23 -

5. Yield per plant

-46.38-93.89 43.07-57.99 -34.09-10.00 17.5-90.0 -10.30-120.71 -12.60-84.10

-96.88-72.45 10.28-21.05 -26.79-7.33 18.0-52.0 -33.69-55.46 -35.13-65.39 -32.71-152.66 9.80-118.80 -19.05-67.78 3.60-56.20 45.50-74.15

-4.49-78.83 9.43-35.85 23.8-87.3 88.92-115.51 -

Nagaraj (1995) Ghosh et al. (1997) Patil (1997) Fageria (2001) Tiwari and Lal (2004) Thakur (2004) Bhutani et al. (1973) Rattan and Saini (1976) Singh et al. (1976) Yoganand (1977) Peter and Rai (1978) Singh et al. (1978) Anbu et al. (1981)

Table 4. Contd..... Characters No. of crosses 30 F1 hybrids 15 F1 hybrids 5 F1 hybrids 5 F1 hybrids 105 F1 parents + 15 parents 28 F1 hybrids 12x12 diallel 28 F1 hybrids 28 F1 hybrids 45 F1 hybrids 12x12 diallel 50 F1 hybrids 6 F1 hybrids 36 F1 hybrids 8x8 half diallel 45 F1 hybrids 91 F1 hybrids 45 F1 hybrids 9 F1 hybrids Range of heterosis (%) over Better parent Commercial check -28.90-69.20 -18.22-73.70 14.60 -41.67-7.68 -166.83-62.36 27.87-72.89 1.19-94.66 -30.47-76.45 13.97 to 94.66 -92.09 to 22.89 30.94 to 53.51 -100.50 to 0.5 17.28 to 48.21 19.1 to 218.5 -37.55-44.98 -45.61-22.07 -30.11 to 47.16 -23.89 to -4.71 19.1 to 105.3 References Rajpur (1987) Kumar et al. (1988) Rammohan (1988) Kanthaswamy and Balakrishnan (1989) Yadav et al. (1989) Dundi (1991) Dod et al. (1992) Singh and Singh (1993) Pujari and Kale (1994) Tendulkar (1994) Dod et al. (1995) Dharmatti (1995) Nagaraj (1995) Ghosh et al. (1997) Patil (1997) Shrivastava et al. (1998) Bhatt et al. (1999) Fageria (2001) Akhilesh and Gulsham (2004)

Mid parent 87.60 -24.96-29.51 -

-25.39-88.85 39.60-14.40 -100.00 to 142.62 -85.92 to 38.11 47.15 to 55.96 -21.97 to 40.27 45.01 to 138.94 72.3 to 373.5

Table 4. Contd..... Characters No. of crosses 8x8 diallel c. Fruit quality parameters 1. Total soluble solids Range of heterosis (%) over Better parent Commercial check -17.41 to 29.89 References Thakur (2004)

Mid parent -

2. Number of locules per fruit

28 F1 hybrids 16 F1 hybrids 5 F1 hybrids 28 F1 hybrids 12x12 diallel 13x13 diallel 45 F1 hybrids 50 F1 hybrids 12x12 diallel 6 F1 hybrids 8x8 half diallel 6x6 diallel 9 F1 hybrid 16 F1 hybrids 12 F1 hybrids 28 F1 hybrids 12x12 diallel 13x13 diallel

222.05 -11.14-2.79 -13.13-55.13 2.76-25.35 -23.78-31.06 -100.00-386.07 -7.16-16.13 -7.95-37.91 -51.35-35.14 77.05 -44.83-37.50 2.00-19.56

148.25 -21.27-5.88 -32.82-65.28 0.00-14.79 -13.41-12.80 -33.94-22.15 -100.00-282.59 0.59-14.79 -3.68-8.85 -57.14-25.00 60.40 -27.97-133.33 -37.5-0.00 -4.50-34.50

1.44-24.22 -7.14-126.19 -40.17-40.88 -100.0-68.11 -17.61-40.88 -18.18-63.64 11.1-35.5 55.10 -45.50-99.82 -

Bhutani et al. (1973) Peter and Rai (1978) Kanthaswamy and Balakrishnan (1989) Dundi (1991) Dod and Kale (1992) Reddy and Reddy (1994) Tendulkar (1994) Dharmatti (1995) Dod et al. (1995) Nagaraj (1995) Patil (1997) Makesh (2002) Tiwari and Lal (2004) Peter and Rai (1978) Anbu et al. (1981) Dundi (1991) Dod and Kale (1992) Reddy and Reddy (1994)

Table 4. Contd..... Characters No. of crosses 9 two parent 42 three parent + 18 four parent 45 F1 hybrids 12x12diallel 50 F1 hybrids 8x8 diallel 36 F1 hybrids 45 F1 hybrids 28 F1 hybrids 13x13 diallel 45 F1 hybrids 16 F1 hybrids 28 F1 hybrids 12x12 diallel 45 F1 hybrids 12x12 diallel 36 F1 hybrids 8x8 half diallel 6x6 diallel 9 F1 hybrids Range of heterosis (%) over Better parent Commercial check -29.54-19.00 References Sundaram et al. (1994)

Mid parent 32.13-11.71

3. Fruit shape index (FSI)

4. Pericarp thickness

-44.88-19.28 -100.00-101.23 -12.22-14.47 46.89-60.45 97.62-141.71 3.66 -35.24-18.58 -12.63-3.63 -13.16-42.73 -17.51-43.03 -7.57-73.29 -11.50-50.27 48.21-57.72 -4.41-14.22 -33.33-55.56 -12.6-41.7

-28.10-36.25 -37.5-21.95 -100.00-87.36 37.88-46.59 -3.18 -50.57-10.12 -27.27-9.34 -55.15-42.42 -22.20-49.47 -17.22-63.16 0.00-9.91 -13.67-40.98 -7.41-5.69 38.63-47.61 -42.89-40.00 -29.4-23.4

-36.10-68.37 -100-103.50 -18.98-8.48 -34.24-36.29 -26.60-47.91 27.66 -44.40-55.5 -39.60-0.77 -18.98-8.48 -40.00-40.00 -30.2-31.4

Tendulkar (1994) Dod et al. (1995) Dharmatti (1995) Patil (1997) Ghosh et al. (1997) Shrivastva et al. (1998) Ashwathappa (1980) Patil (1984) Dundi (1991) Reddy and Reddy (1994) Tendulkar (1994) Peter and Rai (1978) Dundi (1991) Dod and Kale (1992) Tendulkar (1994) Dod et al. (1995) Ghosh et al. (1997) Patil (1997) Makesh (2002) Tiwari and Lal (2004)

Table 4. Contd.....

Range of heterosis (%) over Characters No. of crosses Mid parent 5. Ascorbic acid (mg/100 g) 20 F1s (LxT) 6x6 diallel 9 F1 hybrids 6. Acidity (%) 20 F1 s (LxT) 6x6 diallel -28.7-59.35 -7.82-13.31 46.7-48.9 -17.4-52.25 -20.02-34.34 Better parent -56.04-41.91 -20.40-11.22 22.4-45.4 -37.20-58.12 -31.21-17.37 Commercial check -16.24-15.62 22.4-65.9 -44.13-1.89 Patil and Patil (1988) Makesh et al. (2002) Tiwari and Lal (2004) Patil and Patil (1988) Makesh et al. (2002) References

2.2 DIALLEL MATING

Pushing the tomato crop (Lycopersicon esculentum Mill.) having unfounded superstitions (as unsafety food) that persisted widely to a level of dietary staple is because of the chain of events initiated by tomato workers. With efflux of time, though several cultivars have been developed, seldom serve quantitative and qualitative requirements of farmers. In India, a beginning has been made in this specific use. However, increasing the productivity and production is the need of the hour considering the overall spectrum of situations moving in this direction, the present chapter has been penned in order to capitalise on the literature and achievements available.

2.2.1 Heterosis

The impulse of progress in crop improvement through plant breeding was propelled by a better understanding and an appropriate exploitation of heterosis, the classical term coined by Shull (1914) implying the gain in vigour on crossing two inbreds. A considerable degree of heterosis has been documented and utilized in tomato for various characters ever since the first official report by Hedrick and Booth (1907). Tomato is a self pollinated crop where degree of heterosis was theoretically considered less (Gallias, 1988). However, the unusual high heterosis observed in tomato crop has been attributed to the fact that tomato was basically a highly out crossing genus which was later evolved into a self pollinated crop (Rick, 1965). Based on reports of various scientists, the heterosis of some traits is presented in Table 4.

2.2.2 Combining ability

Sprague and Tatum (1942) developed the concept of combining ability and coined the terms : i) general combining ability (GCA) and ii) specific combining ability (SCA). In pursuit of rendering genetic improvement in crop plants the plant breeders must possess an adequate knowledge of combining ability and allied genetic parameters. Such a knowledge has been steadily increasing on tomato and in this heading few reports have been summarized in Table 5.

Table 5. Review of literature on combining ability in tomato Sl. No. A. 1. 2. B. 1. Characters Growth parameters Plant height (cm) Number of primary branches Yield and yield components Number of flowers per truss SCA variance > GCA variance Significant GCA and SCA variance SCA variance > GCA variance Sonone et al. (1986), Dharmatti et al. (1995) Dod et al. (1992), Bhatt et al. (2001) Sonone et al. (1986), Dharmatti et al. (1995) Dod et al. (1992), Bhatt et al. (2001) Patil (1984), Peter and Rai (1980), Prabhushankar (1990), Bhatt et al. (2001), Cheema et al. (2003) Prabhushankar (1990), Patil (1997) Singh and Mital (1978) Peter and Rai (1980), Bhatt et al. (2001) Dundi (1991), Patil (1984), Dharmatti (1995), Patil (1997) Patil (1997), Dharmatti (1995) Dundi (1991), Nandpuri and Tyagi (1976), Prabhushankar (1990), Swamy and Mathei (1982), Dharmatti (1995), Srivastava et al. (1998) Sonone et al. (1986), Lonkar and Borikar (1988), Singh and Singh (1980), Swamy and Mathai (1982) Combining ability Reference

SCA variance > GCA variance Significant GCA and SCA variance Significant GCA variance Significant GCA and SCA variance SCA variance > GCA variance Significant gca and sca effects

2.

Number of fruits per truss

3.

Number of fruits per plant

Significant gca and sca effects

Significant GCA variance > SCA variance

Table 5. Contd..... Sl. No. Characters Combining ability High SCA variance SCA variance > GCA variance Significant GCA and SCA variance GCA variance > SCA variance Reference Dundi (1991), Govindarasu et al. (1981), Prbhushankar (1990) Dharmatti (1995), Pradeepkumar et al. (1997), Patil (1997), Dod et al. (1995) Bhatt et al. (2001), Sharma et al. (2002) Sonone et al. (1986), Dod et al. (1995), Pradeepkumar et al. (1997), Patil (1997), Sharma et al. (2002) Dharmatti (1995) Sidhu et al. (1981) Cheema et al. (2003), Sharma et al. (2002) Dundi (1991), Patil and Bojappa (1986), Prbhushankar (1990), Swamy and Mathai (1982), Bhatt et al. (2001), Sharma et al. (2002) Lonkar and Borikar (1998), Sharma et al. (1996), Sidhu et al. (1981) Anbu et al. (1980), Lonkar and Borikar (1988), Sharma et al. (1996), Sidhu et al. (1981) Patil (1997) Dundi (1991), Prbahushankar (1990), Swamy and Mathai (1982), Patil (1997), Dharmatti (1995), Srivastava et al. (1998), Cheema et al. (2003)

4.

Average fruit weight

5.

Yield per plant

SCA variance > GCA variance Significance GCA variance > SCA variance Highly significant gca and sca effects Significant GCA and SCA variance

High GCA variance High SCA variance GCA variance > SCA variance Significant gca and sca effects

Table 5. Contd..... Sl. No. Characters Combining ability Significant SCA variance > GCA variance C. 1. Fruit quality parameters Total soluble solids (TSS) Significant ­ SCA variance > GCA variance SCA variance > GCA variance GCA variance > SCA variance Significant gca and sca effect 2. Number of locules per fruit Significant GCA and SCA variance Significant GCA variance > SCA variance SCA variance > GCA variance Significant gca and sca effects Significant SCA variance > GCA variance Dundi (1991), Prabhushankar (1990) Anbu et al. (1980), Govindarasu et al. (1981), Patil (1984), Patil and Bojappa (1986), Dharmatti (1995) Sonone et al. (1986), Sharma et al. (1996), Patil (1997), Dod et al. (1995), Shrivastava (1998) Pradeepkumar (1997), Dhaliwal et al. (1999), Cheema et al. (2003) Peter and Rai (1980), Tarrera and Nuez (1983), Patil (1997) Dundi (1991), Prabhushankar (1990), Singh and Singh (1980) Anbu et al. (1980), Govindarasu et al. (1981), Patil (1984), Sonone et al. (1986) Shrivastava et al. (1998), Dharmatti (1995), Dhaliwal et al. (1999), Cheema et al. (2003) Pradeep kumar et al. (1997) Reference Sonone et al. (1986), Pradeepkumar et al. (1997), Dod et al. (1995), Sajjan (2001), Kulkarni (1999)

Table 5. Contd.....

Sl. No. 3. 4.

Characters

Combining ability GCA variance > SCA variance Dod et al. (1995)

Reference

Fruit shape index (FSI) Pericarp thickness

GCA variance > SCA variance Significant gca and sca effects Significant GCA variance > SCA variance High sca effects Significant sca effects Significant SCA variance > GCA variance Significant gca and sca effects GCA variance > SCA variance

Singh and Mital (1978) Tendulkar (1994), Dundi (1991) Dixit et al. (1980), Rai et al. (1997), Dundi (1991), Prabhushankar (1990), Sharma et al. (1996) Patil and Bojappa (1986) Dundi (1991) Shrivastava et al. (1998) Dhaliwal et al. (1999) Patil (1997), Dod et al. (1995) Patil and Bojappa (1986) Patil and Bojappa (1986) Shrivastava (1998)

5. 6. 7.

Ascorbic acid Acidity Reducing sugar

Significant SCA variance > GCA variance Significant SCA variance > GCA variance Significant GCA variance > SCA variance

III. MATERIAL AND METHODS

The present investigation on variability, heterosis and combining ability in tomato was conducted during 2005-06 at Botany Garden, Department of Genetics and Plant breeding, University of Agricultural Sciences, Dharwad. The details of the materials and methodologies used and the experimental techniques employed for the studies are outlined in this chapter under the following headings. 3.1 Experiment I: Biparental mating 3.2 Experiment II: Diallel mating

3.1 EXPERIMENT I: BIPARENTAL MATING

3.1.1 Location of experimental site and climate

The field experiment was conducted in the Botany garden of Agriculture College, Dharwad during 2005-06. Geographically Dharwad is situated at 15° 26N latitude and 70° 26E longitude at an altitutde of 678 m above mean sea level. The average annual rainfall is about 700-800 mm which is distributed evenly during the cropping period. The monthly maximum and minimum temperatures during hot months (March-April) and during cold months (December-January) ranged from 35.7°C to 38.3°C and 29.17°C to 30.50°C respectively.

3.1.2 Experimental material

The experimental material for the present investigation comprised of following. F1s : Commercial hybrids, MHTM-256 and S-4-14 F3 : S-4-14 Biparental progenies (BIPs) A. Intramating population i. Population M ­ Derived by intermating selected segregants in F2 of MHTM-256 ii. Population S ­ Derived by intermating selected segregants in F2 of S-4-14. B. Intermating population iii. Population M/S ­ Obtained by crossing selected F2 plants in MHTM-256 as female with selected F2 plants of S-4-14. iv. Population S/M ­ Obtained by crossing selected F2 plants in S-4-14 as female with selected F2 plant of MHTM-256.

3.1.3 Experimental methods

The F2 population of MHTM-256 and S-4-14 were sown in nursery during April 2005 and transplanting of seedlings to mainfield was carried out a month after sowing, to effect the crossing to get biparental populations as explained in the sub-topic experimental material. Of the two, the best F2 population (S-4-14) was carried forward to the next generation by selfing and compared with the biparnetal mating population. The seeds of intra mating population in MHTM-256, intra mating population in S-4-14, selfed seeds, inter mating population using MHTM-256 as female and S-4-14 as male and vice versa were sown during November 2005. The seedlings were transplanted to mainfield thirty days after sowing. Five rows each of commercial hybrids MHTM-256 and Namdhari were placed as checks.

3.1.4 Collection of data

The following observations were recorded on all plants in biparental progenies.

3.1.4.1 Plant height Plant height was measured in centimeters from ground level to the tip of the plant at the time of last harvesting. 3.1.4.2 Number of primary branches Number of primary branches per plant was counted at the time of harvest. 3.1.4.3 Number of fruits per plant Number of fruits per plant was counted by pooling the berries from all the pickings. 3.1.4.4 Number of fruits per truss Number of fruits per truss was recorded for the first inflorescence of each plant. 3.1.4.5 Average fruit weight (g) Average fruit weight was computed by using following formula.

Total fruit weight of each plant Average fruit weight (g) = ------------------------------------------ Total number of fruits from each plant 3.1.4.6 Yield per plant (g) Yield was determined by adding the total fruit weight over all pickings from each plant. 3.1.4.7 Fruit shape index Using vernier calipers, the polar (L) and equatorial (D) diameter of fruit was recorded. The shape index was calculated by the ratio (L/D). 3.1.4.8 Number of locules per fruit Number of locules was counted from five fruits taken at random by cutting transversly in the middle. 3.1.4.9 Pericarp thickness (mm) The fruits selected for recording locule number per fruit were sliced at the equatorial plane to measure pericarp thickness in mm. 3.1.5 Statistical analysis The data collected as explained above were subjected to the following statistical analysis. 3.1.5.1 Mean On the basis of individual plant observations, the mean for each character in all the populations was computed as follows.

n

y = 1/n ( yi)

i=1

Where, y = Population mean yi = Individual value n = Number of observations

3.1.5.2 Range The minimum and maximum value on the basis of individual plant observations was used to indicate the range for a given character. 3.1.5.3 Variance In all the populations, variance was computed as follows. Variance = 1/n-1 [(yi - y)²] Where, yi = Individual value y = Population mean n = Number of observations Standard deviation (SD) = Variance = d²/n Where, d = Deviation of individual value from mean n = Number of observations

SD Standard error = ---- n 3.1.5.4 Estimation of genetic parameters Genetic parameters were estimated in the segregating populations viz., inter population mating and intra population mating. Genotypic and phenotypic variance Genotypic and phenotypic variances were computed as follows. Variance of segregating populations viz., F3 as well as BIP population is supposed to have genetic variance (Vg) and environmental variance (Ve) as the main components. VF3/VBIP = Vg + Ve On the contrary, variation in non-segregating generation i.e. F1 is supposed to be only due to the environment and equated to Ve. Ve = VF1 Therefore, VgBIP = VBIP ­ Ve Genotypic and phenotypic coefficient of variation The genotypic and phenotypic coefficient of variation was computed according to Burton and Devane (1953). Vg Genotypic coefficient of variation (GCV) = -------- x Vp Phenotypic coefficient of variation (PCV) = -------- x Where, x 100 x 100

Vg = Genotypic variance Vp = Phenotypic variance x = General mean of the character PCV and GCV were categorized as low, moderate and high by following Sivasubramanian and Menon (1973). It is as follows 0 ­ 10 % 10 - 20% 20% and above : Heritability (h²) Heritability in broadsense which is the heritable variation was estimated as the ratio of genotypic variance to the phenotypic variance and expressed in percentage (Hanson et al., 1956). Vg Heritability (h²) = -------- Vp Where, Vg = Genotypic variance Vp = Phenotypic variance Genetic advance (GA) The extent of genetic advance to be expected from selecting five per cent of the superior progeny was calculated by using the following formula given by Robinson et al. (1949). GA = i p h² Where, i = Coefficient intensity of selection which is 2.06 at 5 per cent selection x 100 : : High Low Moderate

p = Phenotypic standard deviation h² = heritability in broadsense Genetic advance as per cent of mean GA GA (as % of mean) = -------- X Where, GA = Genetic advance X = General mean of the character 3.1.5.5 Inter character correlation In all the generations, the simple correlation coefficients were calculated to determine the degree of association of different characters with fruit yield and also among yield components in each of the populations separately. Correlation coefficients were compared against table `r' values (Fisher and Yates, 1963) at (n-2) degrees of freedom at the probability levels 0.05 and 0.01 to test their significance. Simple correlation was computed by using the formula give by Weber and Moorthy (1952) as given below. x 100

Cov xy r= -------- Vx Vy Where, Cov xy = Covariance between the characters x and y Vx = Variance of the character x Vy = Variance of the character y 3.1.5.6 Path coefficient analysis In all the segregating populations path coefficient analysis was carried out using the simple correlation coefficient to know the direct and indirect effects of the yield components on seed yield as suggested by Wright (1921) and illustrated by Dewey and Lu (1957). Standard path coefficients which are the standardized partial regression coefficients were obtained by solving the following set of `P' simultaneous equation through the use of "Doolittle Technique" as described by Goulden (1959). P01 + P02r12 + . . . . . . + Popr1p = r01 P01 + r12 P02 + . . . . . . + r2pPop = r02 P01 + r1p + P02 r2p + . . . . . . + Pop = r0p Where, Po1, Po2 . . . Pop = Direct path coefficient of variables r12, r13 . . . r1p . . rp (p-1) = Possible correlation coefficient between various independent variables r01, r01 . . . r0p = The correlation between dependent variable and independent variables The direct effect of ith variable via jth variable was worked out as (Poj x rij). From simultaneous equations, it is clear that the correlation coefficient is the sum of direct and indirect effects on dependent character. Residual effect (Pox²) was calculated as under. P²ox = 1-(P²01 + 2 P01 P02 r12 + 2 P01 P03 r13 . . 2 P02 P03 r23 + . . + P²op

3.1.6 Superior segregants

In all the BIP and F3 populations separately the number of plants which exceed the performance of the check was (MHTM-256) noted as superior segregants for the fruit yield trait.

3.2 EXPERIMENTAL II: DIALLEL MATING

3.2.1 Location of experimental site and climate

The location of experimental site and climate was explained in the subtopic location of experimental site and climate of biparental mating.

3.2.2 Experimental materials

The experimental material consisted of 7 parents viz., S-22, L-15, CO-3, Sivap, Solan Vajra, Local type and PKM-1. Diallel mating system was utilized for developing hybrids. The salient features of the parental lines are presented in Table 6.

3.2.3 Experimental methods

3.2.3.1 Hybridization programme The parental seeds were sown in nursery beds during kharif 2005. The healthy thirty day old seedlings were used for field planting. For making diallel crosses, unpaired parents planting arrangement was followed.

Table 6. Salient features of parents used in diallel mating

Origin UAS, Dharwad L-15

Parents

Description Cross between NTDR-1 and AVRDC breeding line. Bacterial wilt resistant, medium sized fruits. Capacity to set fruit under high temperature A mutant of CO-1, plant erect, determinate, rich in vitamin C, fruits red, medium size Small fruits, determinate, high TSS Medium size fruits, dark red colour, high ascorbic acid Medium sized fruits of 70 g. Bears about 4 fruits per inflorescence. Longest harvest duration, yields about 1.3 kgs per plant Medium size fruits, average fruit weight of 40g, high yielder Medium size fruits, average fruit weight of 50g, high yielder, dark red colour

TNAU, Coimbatore

CO-3

Local collection, Dharwad TNAU, Coimbatore YSP University of Horticulture and Forestry, Nauni-Solan IIHR, Bangalore YSP University of Horticulture and Forestry, Nauni-Solan

Local type PKM-1 Solan Vajra

S-22 (Arka Vikas) Sivap

For hybridization the floral buds of the female parents were emasculated a day before their opening between 3 to 6 pm with the aid of pointed forceps and bagged to prevent cross pollination. Following morning, pollen grains were collected from freshly opened flowers of the desired male parent and gently applied to the stigma of the emasculated flower. The crossed flowers were tagged for easy identification and bagged just for two days. Seeds were extracted from red ripe fruits. To ensure selfing of parental genotypes, flowers destined to open next day were covered with butter paper bags, then seeds were extracted from such fruits. 3.2.3.2 Evaluation of F1 hybrids Nursery raising Raised nursery beds of size 3.0 m x 1.2 m and 15 cm in height were prepared and sterilized. Each bed was applied with 500 g of 15:15:15 NPK complex fertilizer and mixed thoroughly. The seeds of 7 parents, 21 hybrids and 1 check were sown in rows spaced at 10 cm apart during the first fortnight of November 2005. The beds were regularly watered. To control damping off appropriate control measures were undertaken. Four week old seedlings were transplanted during first fortnight of December in experimental block with 29 entries in randomized block design with 3 replications. Each entry was represented by a single row of 8 plants with a spacing of 60 x 45 cm. The plants were supplied with 115 kg of N, 100 kg of P2O5 and 60 kg of K2O per hectare and irrigation was given as and when needed. Plant protection measures were undertaken during necessity.

3.2.4 Observations recorded

Five random plants from each entry row in each replication were tagged for recording the following observations. The method followed for recording observations for the characters viz., plant height, number of primary branches, number of fruits per plant, number of fruits per truss, number of fruits per plant, average fruit weight, yield per plant, fruit shape index, number of locules per fruit and pericarp thickness was same as that of biparental mating experiment. The other characters for which observations were recorded is explained here below. 3.2.4.1 Number of flowers per truss Number of flowers per truss was recorded for the first inflorescence of each plant and the mean computed. 3.2.4.2 Total soluble solids (TSS) A drop of juice was used to record the TSS (%) with the help of Erma hand refractometer at ambient temperature. 3.2.4.3 Total titrable acidity Citric acid A known volume of sample was diluted with a known volume of distilled water. An aliquot taken from this sample was titrated with 0.1 N NaOH using phenolphthalein as indicator (Anon., 1984).

Titre x Normality x M.eq.wt. of acid % acid = -------------------------------------- Weight of volume of sample Milli equivalent weight of citric acid = 0.06404 x 100

3.2.4.4 Ascorbic acid content of fruits (Sadasivam and Manickam, 1992) The ascorbic acid content in fruit was estimated by 2, 6-dichlorophenol indophenol visual titration method (AOAC, 1970) and values were expressed in mg per 100 g of fruits. The fruit sample of 2.5 g was taken and grinded in metamorphic acid and acetic acid mixture. Final volume was made up to known volume by adding metamorphic acid and acetic acid mixture. The extract was filtered through filter paper (Whatman No. 41) and the filtrate was titrated against 2,6-dichlorophenol indophenol. The titration value was used to calculate ascorbic acid content and expressed as mg/100 g of fruit sample and mean calculated for each treatment. 3.2.4.5 Reducing sugar (Sadasivam and Manickam, 1992) Estimation of reducing sugar by Dinitro-salicylic acid method employs 3,5dinitrosalicylic acid (DNSA) which is reduced in an alkaline medium to 3-amino, 5 nitrosalicylic acid. DNSA reagent was prepared by dissolving 1 g of 3,5-dinitrosalicylic acid in little amount of 2 N NaOH. Procedure Pipette out into a series of labeled test tubes aliquots from stock standard solution and make up to 1 ml in all the tubes and maintain a blank. Add 0.5 ml of DNSA reagent to all the tubes, mix well and keep in boiling water bath for 5 minutes. Cool the tubes and make up to a suitable volume. Read the per cent T of the standard and the sample against reagent blank. Calculate mg of reducing sugar present per gram of the sample based on the standard graph. 3.2.4.6 Total sugar Total sugar was estimated by acid hydrolysis of non-reducing sugar. Reagents 1. Sucrose solution: Dissolve 80 mg of sucrose in distilled water and make up to 50 ml in a volumetric flask 2. 0.1 and 1 N HCl and 1 N NaOH 3. Phenolphathalein indicator solution in alcohol Procedure Place in a test tube 0.5 ml of sucrose solution. Add 0.5 ml of water followed by 1 ml of 1 N HCl and place in a water bath at 50°C for 20 minutes. Cool, add a drop of indicator solution and mix well. Add dropwise 1 N HCl till the solution turns pink due to excess alkali. Reneutralise the excess alkali with 0.1 N HCl which is added dropwise till the solution becomes colourless and make up to known volume. Draw suitable aliquots and estimate reducing sugar present in the hydrolysate by DNSA method. 3.2.4.7 Non-reducing sugar Non-reducing sugar was estimated by following formula. Non-reducing sugar = (Total sugar-reducing sugar) 0.95 3.2.4.8 Juice yield (%) The fruits were cut and blended in a pestle and mortar and filtered through a muslin cloth to get the juice. The juice yield was measured and expressed as per cent of total weight of the fruits. 3.2.4.9 Pulp weight (%) The pulp weight was recorded after the fruits were pulped and juice extracted. The remaining pulp was weighed discarding the seeds and expressed in percentage.

3.2.5 Statistical analysis and interpretation of data

3.2.5.1 Analysis of variance The mean values of the genotypes were subjected to analysis of variance (Panse and Sukhatme, 1967). The model of variance table adopted is given below. Source of variation Replications Genotypes Error Total Where, r = Number of replications g = Number of genotypes 3.2.5.2 Heterosis The magnitude of heterosis was estimated in relation to mid parent (MP), better parent (BP) and commercial check (CC) (MHTM-256) as percentage increase or decrease of F1s over the respective parents. Average values over replications were used for estimating the heterosis over mid parent, best parent and the commercial check (economic check) using the methods of Turner (1953) and Hayes et al. (1956). F1 ­ MP Heterosis over mid parent (MP) = MP X 100 d.f. (r-1) (g-1) (r-1) (g-1) (rg-1) M.S.S. M1 M2 M3 M1+M2+M3 Expected M.S.S. ²e+²g ²e M2/M3 Cal F

P1 + P2 where mid parent = 2

F1 ­ BP Heterosis over better parent (BP) = BP X 100

F1 ­ CC Heterosis over commercial check (CC) = CC Where, F1, BP and CC are mean values of F1 hybrids, better parent and commercial check respectively. The significance of F1 heterosis values was tested by comparing them with CD values obtained separately for MP, BP and CC employing the formula given below. 3/2 x MSSe CD for MP = r X `t' value X 100

2 x MSSe CD for BP and CC = Where, MSSe = Error mean sum of squares r = Number of replications t = Table `t' value at error degrees of freedom 3.2.5.3 Combining ability analysis The variation among the hybrids was further partitioned into genetic components attributed to general combining ability (gca) variances and specific combining ability (sca) variances and effects were analysed by adopting Model-I, method-2 of Griffing's (1956), since the present study includes parents and F1s (without reciprocals). The statistical procedure assumes the following mathematical model. xij = µ + gi + gj+ sij + 1/bc eijkl

k 1

r

X `t' value

Where, ij = 1, 2, . . . . n k = 1, 2, . . . . b l = 1, 2, . . . . c µ = Population mean gi = General combining ability (gca) effect of ith parent gj = General combining ability (gca) effect of j parent eijkl = Environmentla effect associated with the ijklth individual observation n = Number of parents b = Number of replications c = Number of individual in each replication The model assumes that a = igi = 0 and Sij = 0 (for each i) b = The error (eijkl) is normally and independently distributed with mean equal to zero and variance equals ²c. 3.2.5.4 Combining ability variance The analysis of variance table for combining ability is as follows Source GCA SCA Error Where, Sg = 1/(n+2) [i(xi + xij)²] 4/n x² Ss = x²ij 1/(n+2) (xi+xij)² + 2/(n+1) (n+2)

i j th

d.f. (n-1) n(n-1)/2 M

SS Sg Ss Se

MSS Mg Ms Me

EMSS ²+²s+(n+2)²g ²+²s ²e

x².........

xi = Total of i row xii = Value of i parent x.. = Grand total of the diallel table xij = Value of the cross between i and j parents N = Number of parents The estimation of the Me' the error term in the above table was obtained as Me' = Me/b, where Me is the error variance estimated in the analysis of variance of the experiment and `b' is the number of replications. The general and specific combining ability variances were calculated based on the mean sum of squares as follows. Where, (Mg-Me) (n-1) ²g = (n+2) (Mg-Me) (n-1) ²p = 2 The significance was tested by the `F' ratio against M1e. 3.2.5.5 Estimation of combining ability effects The general combining ability (gca) effects and specific combining ability (sca) effects were estimated as follows. 1 gca effects = gi = n+2 [(xi+xij)- 2/n x..]

th th th

th

1 sca effects =sij = Xij n+2 Where, xj = Total of j column xjj =Value of jth parent Remaining n, xii, x . .. are same as mentioned above 3.2.5.6 Test of significance for the combing ability effects

th

2 [xi+xii+xj+xjj) + ------------x . . . (n+1) (n+2)

Standard error (SE) for different estimates was obtained from the following formulae. SE (gi)= [(n-1)²e/n (n+2)]½ SE (gi-gj)= [2²(n+2)]½ n(n-1)²e SE (Sij)= ------------ (n+1) (n+2) Critical difference (CD) for testing the significance of the combining ability estimates was obtained by multiplying the respective standard error with table `t' value at error degrees of freedom.

IV. EXPERIMENTAL RESULTS

The presence of genetic variability for economic traits is a key for improving the varieties. This investigation was carried out to assess and compare the variability generated by different biparental mating schemes and selfing using two commercial hybrids MHTM-256 and S-4-14. The data obtained were subjected to statistical analysis to find out mean, range, genetic parameters and correlation among the characters under study. Another experiment with 21 hybrids, seven parents and one commercial check hybrid (MHTM-256) was carried out to determine heterosis for yield and yield components and also to estimate the combining ability effects and variances for yield and yield components. The results of the present investigations are presented under the following captions. 4.1 Experiment I: Biparental mating 4.2 Experiment II: Diallel mating

4.1 EXPERIMENT I: BIPARENTAL MATING

The results of the present investigation are presented under the following captions. 4.1.1 Mean 4.1.2 Range 4.1.3 Variance 4.1.4 Heritability and genetic advance 4.1.5 Inter character correlation 4.1.6 Path coefficient analysis 4.1.7 Identification of superior segregants

4.1.1 Mean

The mean values of different characters in segregating populations are presented in Table 7 and the results are explained here under. In general the intra mating populations (Population M and population S) showed higher mean values for the characters viz, plant height (45.63 and 53.11 cm), primary branches (3.97 and 4.36), fruits per plant (13.05 and 11.92), average fruit weight (34.70 and 44.30 g), fruit yield (473.42 and 542.80 g), locules per fruit (4.10 and 4.36) and fruit shape index (1.12 and 1.03). Where as the inter mating populations (population M/S and population S/M) showed higher means for the character, fruits per truss. However, there was no difference among the inter and intra mating populations in general for the mean of the trait pericarp thickness though population S recorded highest mean for this trait (0.39 cm). Compared to BIPs in general, F3 showed low mean values for characters viz., primary branches (3.3), fruits per truss (2.06), average fruit weight (40.00 g), fruit shape index (0.98) and locules (3.90), but showed relatively high values for characters plant height (50.90 cm), pericarp thickness (0.38) and yield per plant (521.30 g). Whereas for character fruits per plant, the F3 mean value (12.53) was comparable with BIPs. As compared to checks (MHTM-256 and Namdhari hybrid) the BIPs exhibited lower mean values for the traits plant height, fruits per truss, fruits per plant and fruit yield per plant since range for these characters and also for other characters understudy was higher in case of the BIPs.

Table 7. Mean performance of F3 population, check and BIP populations in respect of nine characters in tomato Character Populations Plant height (cm) 45.63 53.11 46.46 49.04 50.90 73.30 70.1 Number of primary branches 3.97 4.36 3.58 3.98 3.30 3.66 3.70 Average fruit weight (g) 34.70 44.30 30.80 42.17 40.00 31.80 32.90 Fruit shape index 1.12 1.03 0.84 1.01 0.98 0.91 0.80 Pericarp thickness (cm) 0.3 0.39 0.34 0.34 0.38 0.34 0.36

Fruits per truss

Fruits per plant

Locules per fruit

Fruit yield (g)

Population M Population S Population M/S Population S/M F3 MHTM-256 (F1) (check) Namdhari hybrid

2.41 2.08 2.46 2.30 2.06 3.33 3.20

13.05 11.92 12.69 10.88 12.30 31.3 30.0

4.10 4.36 3.83 3.32 3.90 3.11 3.20

473.42 542.8 401.2 456.6 521.30 973.3 988.4

Table 8. Range for nine traits in F3 population, check and BIP population in tomato Character Populations Plant height (cm) 20.0-90.0 30.0-75.0 20.0-70.0 25.0-105.0 30.0-80.0 69.0-75.0 68.0-74.0 Number of primary branches 2.0-7.0 2.0-7.0 2.0-6.0 2.0-6.0 2.0-6.0 3.0-4.0 3.0-4.0 Average fruit weight (g) 10-55.70 19-67.50 10.0-135.0 10.0-160.0 22.31-90.0 30.3-33.5 31.5-34.0 Fruit shape index 0.5-1.5 0.6-1.5 0.60-1.25 0.28-1.5 0.63-1.26 0.80-0.94 0.75-0.88 Pericarp thickness (cm) 0.2-0.5 0.2-0.5 0.2-0.5 0.2-0.5 0.3-0.5 0.3-0.4 0.3-0.44

Fruits per truss

Fruits per plant

Locules per fruit

Fruit yield (g)

Population M Population S Population M/S Population S/M F3 MHTM-256 (F1) (check) Namdhari hybrid

2.0-4.1 1.2-3.3 1.5-4.5 1.2-4.5 1.5-4.2 2.0-3.0 2.0-3.0

1-47 1-32 3-78 1-52 1-42 29-33 28-32

2.0-7.5 2.5-8.0 2.0-7.0 2.0-8.0 2.0-6.0 3.0-4.0 3.0-4.0

10-1510 30-1530 30-3340 10-1650 60-1650 750-1000 790-1200

4.1.2 Range

The range observed for nine characters in respect of all the populations (Table 8) are presented here under. The characters plant height (20.0 to 70.0 cm and 25.0 to 105.0 cm), fruits per truss (1.5 to 4.5 and 1.2 to 4.5) fruits per plant (3 to 78 and 1 to 52), average fruit weight (10.0 to 135.0 g and 10.0 to 160.0 g), fruit yield (30 to 3340 g and 10 to 1650 g) and fruit shape index (0.60 to 1.25 and 0.28 to 1.50) recorded higher range for the inter mating populations as compared to intra mating populations and the character, primary branches per plant (2.0 to 7.0 for population M/S and population S/M respectively) showed higher range for the intra mating population. The characters pericarp thickness and locules per fruit exhibited quite similar range for both the populations. The range observed in F3 for different characters like plant height (30.0 to 80.0 cm), primary branches (2.0 to 6.0), fruits per truss (1.5 to 4.2), fruit per plant (1 to 42), average fruit weight (22.3 to 90.0 g) locules per fruit (2.0 to 6.0) and fruit shape index (0.63 to 1.26) was narrow than BIPs, but for character fruit yield F3 recorded wider range (60-1650 g) compared to population M and population S.

4.1.3 Variance

The genotypic and phenotypic variance, genotypic and phenotypic co-efficient of variation as observed in all the BIP and F3 populations for all the characters (Table 9) are presented here under. 4.1.3.1 Plant height The genotypic variance (51.30 and 35.384) and phenotypic variance (55.66 and 40.20) was higher for the inter mating populations (Population M/S and population S/M) as compared to intra mating populations. However, the GCV and PCV are quite comparable for inter and intra mating populations. The F3 population showed Vg and Vp (36.44 and 41.00 respectively) quite higher than population S/M and S but less than M/S and M. The GCV and PCV followed the Vp and Vg. 4.1.3.2 Primary branches High Vg of 1.50 and 1.92 was observed in population M and population S respectively (intra mating populations) for the trait under consideration as compared to population M/S (0.89) and population S/M (0.55). The Vp, GCV and PCV followed the similar pattern as that of Vg. F3 population showed Vg and Vp (0.47 and 0.75 respectively) comparable to population S/M but less than rest of BIPs. 4.1.3.3 Fruits per truss The genotypic variance for inter mating population for the trait fruits per truss was quite lower than the intra mating population (0.239 and 0.472) and so also the phenotypic variance. Though GCV was higher in intra mating population (23.50 and 28.50) than inter mating population, the PCV was quite comparable in both the populations. The F3 population showed least Vg (0.052) and Vp (0.282) compared to BIPs. It showed GCV of 11.06 and PCV of 25.70 per cent. 4.1.3.4 Fruits per plant Quite high Vg was observed in case of the intra mating populations (47.18 and 125.39) for the trait fruits per plant as compared to inter mating population (95.35 and 51.36 for population M/S and population S/M respectively). The Vp also had similar pattern as that of Vg. Though the population S showed highest GCV (85.80) and PCV (89.15) for the character under consideration these two parameters in general not differed for the both the populations viz., inter mating population and intra mating population. F3 population showed Vg (72.06) and Vp (82.07) higher than populations S and S/M but lower than population M and M/S. It showed GCV of (82.07) and PCV of (69.00).

Table 9. Estimate of genetic parameters for nine characters in biparental and F3 populations of tomato Sl. No. 1 Character Plant height (cm) Population M Population S Population M/S Population S/M F3 (S 4-14) Population M Population S Population M/S Population S/M F3 (S 4-14) Population M Population S Population M/S Population S/M F3 (S 4-14) Population M Population S Population M/S Population S/M F3 (S 4-14) Population M Population S Population M/S Population S/M F3 (S 4-14) Vg 34.18 38.18 51.30 35.84 36.64 1.50 1.92 0.89 0.55 0.47 0.239 0.472 0.460 0.127 0.052 47.18 125.39 95.35 51.36 72.06 109.92 157.70 184.42 159.90 106.54 Vp 38.54 42.54 55.66 40.20 41.00 1.78 2.20 1.17 0.83 0.75 0.469 0.702 0.692 0.357 0.282 57.19 135.40 105.36 61.37 82.07 116.02 164.60 191.72 160.26 112.84 GCV (%) 11.11 14.14 15.14 12.11 11.89 30.6 38.3 29.2 20.71 23.30 23.50 28.50 27.50 15.49 11.06 57.62 85.80 76.90 65.86 69.00 23.66 36.18 44.09 29.98 25.80 PCV (%) 12.10 17.07 16.16 13.03 12.57 33.3 41.08 33.50 25.40 29.40 22.50 34.76 33.81 25.90 25.70 63.49 89.16 80.80 72.00 73.65 24.30 36.90 44.90 30.01 26.55 h²bs (%) 88.68 89.92 92.16 89.10 83.10 84.20 87.63 76.06 66.26 62.60 50.98 67.23 66.47 35.50 18.43 82.49 92.50 90.41 83.68 87.80 94.56 96.75 96.73 96.06 94.40 GA 11.34 12.08 14.16 11.63 10.96 2.30 2.60 1.60 1.20 1.80 0.72 1.16 0.928 0.44 0.20 6.23 10.76 9.28 1.35 1.63 20.98 25.57 27.59 25.05 20.65 GA (% of mean) 21.35 26.47 30.47 23.71 21.53 57.50 72.00 49.68 50.83 61.22 34.61 48.13 37.70 19.13 9.70 52.26 76.92 70.92 12.40 13.30 47.30 73.68 89.50 59.40 51.62

2

Number of primary branches

3

Fruits per truss

4

Fruits per plant

5

Average fruit weight (g)

Table 9. Contd..... Sl. No. 6 Character Locules per fruit Population M Population S Population M/S Population S/M F3 (S 4-14) Population M Population S Population M/S Population S/M F3 (S 4-14) Population M Population S Population M/S Population S/M F3 (S 4-14) Population M Population S Population M/S Population S/M F3 (S 4-14) Vg 1.47 1.89 0.86 0.52 0.44 0.125 0.123 0.008 0.035 0.0467 0.0049 0.0042 0.0019 0.0036 0.0035 13858.34 16701.91 15341.03 10064.89 15337.73 Vp 1.75 2.17 1.14 0.799 0.718 0.135 0.133 0.018 0.045 0.057 0.0082 0.0076 0.0052 0.0069 0.0068 14869.89 17713.52 16352.58 11076.44 16349.28 GCV (%) 27.80 33.50 24.20 21.70 17.00 34.3 31.3 10.64 18.70 22.05 17.94 20.25 12.82 17.64 15.56 21.68 27.32 30.80 21.97 23.75 PCV (%) 30.34 35.90 27.87 26.92 21.72 35.65 32.56 15.97 21.00 24.36 23.21 27.24 21.20 24.43 21.70 22.46 28.00 31.87 23.04 24.50 h²bs (%) 84.00 87.09 75.43 65.08 61.28 92.54 92.48 44.44 77.70 82.45 59.75 55.26 36.53 52.17 51.47 93.19 94.28 93.81 90.86 93.81 GA 2.289 2.640 1.66 1.20 1.10 0.70 0.69 0.13 0.34 0.41 0.11 0.10 0.054 0.089 0.087 225.99 250.99 239.35 187.70 239.32 GA (% of mean) 52.50 64.30 43.30 36.14 28.20 67.96 61.60 15.47 33.66 41.83 28.20 31.25 15.88 26.17 22.89 41.63 53.01 59.65 41.10 45.90

7

Fruit shape index

8

Pericarp thickness (cm)

9

Yield (g)

35

GCV (%) PCV (%)

30

25

Percentage

20

15

10

5

0

M

S

M/S Population

S/M

F3 (S 4-14)

Fig. 1. Estimate of GCV and PCV (%) for fruit yield in biparental and F3 populations of tomato

Fig. 1. Estimate of GCV and PCV (%) for fruit yield in biparental and F3 populations of tomato

GCV (%)

90

PCV (%)

80 70 60

Percentage

50 40 30 20 10 0

M

S

M/S Population

S/M

F3 (S 4-14)

Fig. 2. Estimate of GCV and PCV (%) for fruits per plant in biparental and F3 populations of tomato

Fig. 2. Estimate of GCV and PCV (%) for fruits per plant in biparental and F3 populations of tomato

4.1.3.5 Average fruit weight The inter mating populations, population M/S and population S/M, showed higher Vg (184.42 and 159.90), Vp (191.72 and 160.26), GCV (44.09 and 29.98) and PCV (44.90 and 30.01) than the intra mating population for the above trait. The least Vg (106.54) and Vp (112.84) was exhibited by selfed progeny compared to BIPs. F3 showed GCV (25.80) and PCV of 26.55. 4.1.3.6 Locules per fruit Intra mating populations showed better genotypic variance (1.47 and 1.89), phenotypic variance (1.75 and 2.17), GCV (27.80 and 33.50) and PCV (30.34 and 35.90) for the trait under study than the inter mating population. The GCV and PCV in F3 was 17.00 and 21.72 respectively lower than any of BIPs. 4.1.3.7 Fruit shape index Compared to inter mating populations the intra mating populations had higher Vg (0.125 and 0.123), Vp (0.135 and 0.133), GCV (34.30 and 31.30) and PCV (35.65 and 32.56). F3 showed Vg (0.0467) and Vp (0.057) slightly higher than population S/M, but less than intra mating population. It showed GCV of 22.05 and PCV of 24.36. 4.1.3.8 Pericarp thickness It was observed that the Vg and Vp was very less in each of the populations under study for the above trait. The Vg (0.0049 and 0.0042), Vp (0.082 and 0.0076), GCV (17.94 and 20.25) and PCV (21.21 and 27.24) was quite high in case of intra mating populations than inter mating populations. F3 showed Vg (0.0035) and Vp (0.0068) comparable to Vg and Vp of population S/M but less than intramating population. It showed GCV of 15.56 and PCV of 21.70. 4.1.3.9 Yield per plant High genotypic variance and phenotypic variance was recorded by intra mating population (13858.34 and 16701.91 for Vg and 14869.89 and 17713.52 for Vp) compared to inter population mating (15341.03 and 10064.89 for Vg and 16352.58 and 11076.44 for Vp). Where as GCV (30.80 and 21.97) and PCV (31.87 and 23.04) was quite higher in inter mating populations than intra mating population. The selfed progeny showed Vg (23.75) and Vp (24.50) higher than population S/M and S but less than rest of the two populations. Though the population S showed higher Vg (16701.91) and Vp (17713.52) for the character under consideration the higher GCV and PCV of 30.80 and 31.87 respectively was recorded by the population M/S.

4.1.4 Heritability and genetic advance

The estimates of heritability and genetic advance in all the segregating populations for all the characters are given in Table 9 and presented here under. 4.1.4.1 Plant height Quite high heritability in broad sense of 92.16 and 89.10 was observed in population M/S and population S/M respectively compared to the intra mating populations. The GA also showed similar pattern as that of heritability in broad sense. Heritability in broad sense was high in BIPs compared to F3 (83.1%). The expected genetic advance and GA as % of mean in F3 was 10.96 and 21.53 respectively which was lower when compared to to BIPs. 4.1.4.2 Primary branches High heritability in broad sense (84.20 and 87.63) and expected genetic advance (2.30 and 2.60) was observed in intra mating populations (population M and S respectively) as compared to the inter population mating. Heritability of 62.60 and expected GA of 1.80 observed for F3 population.

4.1.4.3 Fruits per truss As compared to inter mating population the intra mating population showed quite higher magnitude of heritability in broad sense (50.98 and 67.23) and expected genetic advance (0.72 and 1.16). The GA as per cent of mean followed the GA. The GA as per cent of mean for intra mating populations (34.61 for population M and 48.13 for population S) was higher compared to inter mating populations (37.70 for population M/S and S/M8.13 for population S/M) for this trait. The lowest heritability was observed in selfing progeny (18.43) compared to BIPs. The magnitude of genetic advance was 0.20 in F3 population. GA as per cent mean for F3 population was 9.70. 4.1.4.4 Fruits per plant The heritability in broad sense (82.49 for population M, 92.50 for population S, 90.41 for population M/S and 83.68 for population S/M) was comparable among the BIPs. The GA was observed to be higher in the intra mating population with population S showing highest of 10.76 and lowest for population S/M (1.35). As far as GA as per cent of mean is concerned similar pattern as that of GA was observed with highest of 76.92 showed by population S. F3 showed heritability 87.80 and GA of 1.63. 4.1.4.5 Average fruit weight Maximum heritability was observed for this character compared to any of the characters. In all the populations of BIP the heritability in broad sense was comparable, with highest being 96.75 showed by population M and least being 94.56 showed by population M. The expected genetic advance recorded was 27.59, 25.57, 25.05 and 20.98 by population M/S, population M, population S/M and population S respectively. Similar trend was observed with respect to genetic advance as per cent mean with only change that the higher value was showed by population M/S (89.5) and lowest by population S (41.63). F3 population showed the heritability in broad sense 94.40 and it showed expected genetic advance 20.65. 4.1.4.6 Locules per fruit The heritability in broad sense of 84.00 and 87.09 was recorded in population M and population S respectively which was higher than inter mating population (75.43 and 65.08 for population M/S and population S/M respectively). The expected GA was also higher for intra mating population (2.28 and 2.64 for population M and S respectively) compared to intermating population for this trait. GA as per cent of mean of 52.50 and 64.30 was observed for population M and S respectively which was higher than 43.30 and 36.14 for population M/S and S/M respectively. The least heritability for above character was shown by F3 population (61.28). GA as per cent mean of 1.10 showed by F3 population. 4.1.4.7 Fruit shape index High magnitude of heritability was observed for intra mating populations (92.54 and 92.48) as compared to inter mating populations (44.44 and 77.70). The extent of expected genetic advance was observed to be higher in intra mating population than inter mating population. The GA as per cent of mean of 67.96 and 61.60 was observed in population M and S respectively which was higher than 15.47 and 33.66 observed for population M/S and S/M respectively. F3 showed heritability of 82.45 which is higher than population S/M and population M/S. The GA of 0.41 was showed by F3 population. 4.1.4.8 Pericarp thickness The heritability in broad sense for the intra mating populations (59.75 for population M and 55.26 for population S) was quite higher than inter mating populations (36.53 for population M/S and 52.17 for population S/M). In general the expected genetic advance was less for all of the BIPs but higher for the intra mating populations (0.11 and 0.10 for population M and respectively) compared to inter mating populations (0.054 and 0.089 for population M/S and S/M respectively). The GA as per cent of mean followed similar patter as that of expected GA. Heritability of 51.47 and GA as per cent mean of 22.89 showed by F3 population.

4.1.4.9 Yield per plant Quite high heritability of 93.19 and 94.28 was observed for intra mating populations than for inter mating populations (93.81 and 90.86). The expected GA also showed similar pattern as that of heritability with values being 225.99 and 250.99 for population M and population S respectively. But GA as per cent of mean was higher for inter mating population and population M/S showed highest GA as per cent of mean (59.65) for the trait under consideration. Heritability in broad sense of 93.81 was observations in F3 population. The estimated genetic advance values were 239.35 in F3. It showed GA (45.90) lower than population M/S and M.

4.1.5 Inter character correlations

Analysis of simple correlation coefficients of yield components with fruit yield and among themselves in respect of segregating populations are presented in Tables 10 to 14 and explained here under. Correlation studies in the intra mating populations (population M and population S) revealed that the traits, plant height, number of primary branches, number of fruits per plant, fruits per truss and average fruit weight had positive and significant correlation with the fruit yield per plant. Similarly fruits per truss, plant height, primary branches showed significant and positive correlation with number of fruits per plant. Fruits per truss is positively and significantly correlated with plant height and primary branches, so also the primary branches per plant positively and significantly correlated with plant height and average fruit weight. It was also observed that plant height and primary branches, locules per fruit and pericarp thickness significantly and positively correlated. Pericarp thickness showed significant association with fruit yield in population S, changed to non significant in population M. The average fruit weight which was negatively associated with number of fruits per plant in population S changed into positive association in population M. The association of fruit yield per plant with number of fruits per plant, plant height, and primary branches was positive and significant in inter mating population (population M/S and population S/M). It was also observed that the traits fruits per truss, plant height and primary branches were positively and significantly correlated with number of fruits per plant. Further the traits plant height and primary branches were positively and significantly correlated with fruits per truss but average fruit weight was negatively and significantly correlated with fruits per truss. The association between fruit yield per plant with fruits per truss and average fruit weight which was non significant and positive in population S/M turned out to be significant and positive in population M/S. Similarly, the association between average fruit weight with plant height, primary branches and pericarp thickness which was non-significant in population S/M was turned to be significant in population M/S. In F3 population yield per plant showed positive significant association with number of fruits per plant, average fruit weight, plant height and primary branches. Plant height showed positive significant association with primary branches and number of fruits per plant. The primary branches showed positive association with number of fruits per plant. The pericarp thickness exhibited positive and significant association with locules per fruit.

4.1.6 Path co-efficient analysis

The simple correlation coefficients of different characters with fruit yield under study were subjected to path coefficient analysis for estimating the direct and indirect effects of component traits separately on fruit yield which was considered as dependent variable. The direct and indirect effects of various component traits on fruit yield in respect of different BIPs and F3 population are given in Tables 15 to 19 and explained separately here under. 4.1.6.1 Intra mating populations Plant height exhibited negative direct effect (-0.186 for population M and ­0.0514 for population S) on fruit yield while though it had positive and significant association with fruit yield (0.79 and 0.678 for population M and S respectively) in intra mating population. The indirect effect of plant height via other characters except primary branches (0.581 and 0.531 for population M and S respectively) and fruits per plant (0.1385 and 0.1964 respectively for population M and S) was negligible.

Table 10. Nature of association among nine traits in biparental (BIP) Population M of tomato Character Plant height 1.000 Number of primary branches 0.772** 1.000 Fruits per truss 0.680** 0.652** 1.000 Fruits per plant 0.767** 0.756** 0.754** 1.000 Average fruit weight 0.396** 0.262* -0.116 0.108 1.000 Locules per fruit 0.035 0.193 0.101 -0.054 0.491** 1.000 Fruit shape index 0.137 -0.195 -0.124 -0.076 -0.014 -0.032 1.000 Pericarp thickness 0.064 0.081 0.075 0.122 0.410** 0.05 0.126 1.000 Fruit yield

Plant height No. of primary branches Fruits per truss Fruits per plant Average fruit weight Locules per fruit Fruit shape index Pericarp thickness Fruit yield

0.798** 0.749** 0.795** 0.905** 0.406** 0.049 -0.128 0.126 1.000

Table 11. Nature of association among nine traits in biparental (BIP) Population S of tomato Character Plant height 1.000 Number of primary branches 0.586** 1.000 Fruits per truss 0.503** 0.636** 1.000 Fruits per plant 0.588** 0.689** 0.600** 1.000 Average fruit weight 0.691** 0.364** 0.107 -0.245 1.000 Locules per fruit 0.119 0.176 0.066 0.252 0.466** 1.000 Fruit shape index 0.125 0.154 0.188 0.163 -0.067 -0.094 1.000 Pericarp thickness 0.059 0.140 -0.02 0.167 0.563** 0.141 -0.264 1.000 Fruit yield

Plant height No. of primary branches Fruits per truss Fruits per plant Average fruit weight Locules per fruit Fruit shape index Pericarp thickness Fruit yield

0.678** 0.711** 0.530** 0.948** 0.494** 0.137 0.164 0.287* 1.000

Table 12. Nature of association among nine traits in biparental (BIP) Population M/S of tomato Character Plant height 1.000 Number of primary branches 0.626** 1.000 Fruits per truss 0.492** 0.458** 1.000 Fruits per plant 0.690** 0.543** 0.492** 1.000 Average fruit weight 0.204** 0.174* -0.218** 0.075 1.000 Locules per fruit 0.022 0.049 0.037 0.133 0.197* 1.000 Fruit shape index 0.270 -0.102 -0.075 0.004 -0.094 -0.062 1.000 Pericarp thickness 0.018 0.104 0.015 0.112 0.221** 0.111 -0.076 1.000 Fruit yield

Plant height No. of primary branches Fruits per truss Fruits per plant Average fruit weight Locules per fruit Fruit shape index Pericarp thickness Fruit yield

0.702** 0.563** 0.430** 0.939** 0.298** 0.1005 -0.066 0.1001 1.000

Table 13. Nature of association among nine traits in biparental (BIP) Population S/M of tomato Character Plant height 1.000 Number of primary branches 0.581** 1.000 Fruits per truss 0.254** 0.272** 1.000 Fruits per plant 0.664** 0.628** 0.388** 1.000 Average fruit weight 0.119 0.106 -0.200** -0.026 1.000 Locules per fruit 0.06 0.044 0.019 0.023 0.190 1.000 Fruit shape index 0.085 0.067 0.018 0.077 -0.032 -0.129 1.000 Pericarp thickness 0.179 0.058 -0.016 0.067 0.132 0.105 -0.011 1.000 Fruit yield

Plant height No. of primary branches Fruits per truss Fruits per plant Average fruit weight Locules per fruit Fruit shape index Pericarp thickness Fruit yield

0.655** 0.664** 0.154 0.912** 0.155 0.092 0.081 0.137 1.000

Table 14. Nature of association among yield and different characters in F3 Population of tomato Character Plant height 1.000 Number of primary branches 0.516** 1.000 Fruits per truss 0.158 0.218 1.000 Fruits per plant 0.562** 0.639** 0.324** 1.000 Average fruit weight 0.017 0.027 -0.257* -0.133 1.000 Locules per fruit 0.0045 -0.055 -0.107 -0.016 0.416** 1.000 Fruit shape index 0.171 0.066 0.010 0.142 0.114 -0.180 1.000 Pericarp thickness 0.035 -0.085 -0.06 -0.172 0.162 0.202* 0.007 1.000 Fruit yield

Plant height No. of primary branches Fruits per truss Fruits per plant Average fruit weight Locules per fruit Fruit shape index Pericarp thickness Fruit yield

0.558** 0.654** 0.104 0.938** 0.216* 0.145 0.170 -0.088 1.000

Table 15. Path coefficient analysis of different characters towards fruit yield per plant in BIP Population M of tomato Character Plant height -0.01860 0.09170 -0.01260 0.09620 0.08760 0.03289 -0.04950 0.10040 Number of primary branches 0.58180 0.04870 0.11320 0.04070 0.04100 0.01415 -0.01430 0.03611 Fruits per truss 0.08680 0.08330 0.12800 -0.01420 -0.00730 0.02850 0.00346 0.01330 Fruits per plant 0.13850 0.57300 0.57300 0.75900 0.01267 -0.00063 0.00253 -0.00480 Average fruit weight 0.03760 -0.01435 0.03170 0.03670 0.35300 0.00924 -0.00940 0.15069 Locules per fruit -0.00110 -0.00620 -0.00360 0.00140 -0.01657 -0.03290 0.00007 -0.01710 Fruit shape index 0.00276 0.00391 0.00220 0.00132 -0.00020 0.00004 -0.02030 -0.00330 Pericarp thickness -0.03340 -0.03550 -0.03660 -0.01700 -0.05500 -0.06770 -0.02090 -0.13000 Correlation with fruit yield 0.790** 0.740** 0.795** 0.905** 0.406** 0.049 -0.128 0.127

Plant height No. of primary branches Fruits per truss Fruits per plant Average fruit weight Locules per fruit Fruit shape index Pericarp thickness

Diagonal values represent direct effects *Significant at 5% level of probability **Significant at 1% level of probability

Table 16. Path coefficient analysis of different characters towards fruit yield per plant in BIP Population S of tomato Character Plant height -0.05140 0.09110 -0.02500 -0.05830 0.16045 0.04140 0.13117 0.02220 Number of primary branches 0.53170 0.05970 0.02520 0.05090 -0.00800 -0.00860 -0.01490 0.00619 Fruits per truss -0.04540 -0.05900 0.05349 -0.02920 -0.03400 0.11493 -0.00690 0.01010 Fruits per plant 0.19640 0.64810 0.58270 0.93450 0.01830 -0.02110 -0.00660 -0.02080 Average fruit weight 0.03460 -0.02980 0.03820 0.04140 0.29640 0.01120 0.00843 0.10189 Locules per fruit 0.01120 0.00517 0.02250 0.00740 0.01060 0.02750 -0.01110 0.00948 Fruit shape index 0.00530 0.00586 0.02660 0.00583 -0.00100 -0.01680 0.04150 -0.00976 Pericarp thickness -0.01280 -0.00540 0.00210 -0.00400 -0.01850 -0.01090 0.00745 -0.03160 Correlation with fruit yield 0.6785** 0.7119** 0.5307** 0.9488** 0.4948** 0.1375 0.1643 0.2877*

Plant height No. of primary branches Fruits per truss Fruits per plant Average fruit weight Locules per fruit Fruit shape index Pericarp thickness

Diagonal values represent direct effects *Significant at 5% level of probability **Significant at 1% level of probability

Table 17. Path coefficient analysis of different characters towards fruit yield per plant in BIP Population M/S of tomato Character Plant height 0.04560 0.03150 -0.02250 -0.01600 0.06610 0.05206 -0.00280 0.00570 Number of primary branches 0.61388 0.05975 0.02668 0.01699 -0.00440 -0.00140 0.00250 -0.00820 Fruits per truss -0.01850 -0.01730 0.03763 0.03147 0.00900 0.02710 -0.06790 0.04998 Fruits per plant 0.04660 0.48330 0.37700 0.88960 0.00562 0.01470 -0.01230 0.00990 Average fruit weight 0.02035 0.02855 0.01490 0.01766 0.22868 0.00811 -0.00460 0.05940 Locules per fruit 0.00191 0.00147 0.00022 0.00079 0.00070 0.00593 -0.00160 0.00077 Fruit shape index -0.00440 -0.00230 -0.00120 -0.00005 -0.00480 -0.00420 0.01600 -0.00440 Pericarp thickness -0.00290 -0.00230 -0.00290 -0.00150 -0.00288 -0.00172 0.00360 -0.01320 Correlation with fruit yield 0.70268** 0.563** 0.43027** 0.93928** 0.29886** 0.10057 -0.0666 0.10012

Plant height No. of primary branches Fruits per truss Fruits per plant Average fruit weight Locules per fruit Fruit shape index Pericarp thickness

Diagonal values represent direct effects *Significant at 5% level of probability **Significant at 1% level of probability

Table 18. Path coefficient analysis of different characters towards fruit yield per plant in BIP Population S/M of tomato Character Plant height 0.00043 0.03976 0.00001 -0.00090 -0.12300 0.02072 0.06788 0.00456 Number of primary branches 0.57590 0.07690 0.00900 0.00600 0.00001 -0.00005 -0.00005 0.00002 Fruits per truss -0.00060 -0.00070 0.02380 0.00003 0.00001 0.05157 -0.00780 0.03588 Fruits per plant 0.03260 0.54510 0.10200 0.86600 0.01180 0.00003 0.00004 0.00000 Average fruit weight 0.04490 0.00250 0.02120 0.04844 0.25926 0.00340 0.00530 0.06183 Locules per fruit 0.00088 0.00064 0.00028 0.00035 0.00290 0.01451 -0.00190 0.00211 Fruit shape index 0.00171 0.00134 0.00039 0.00152 -0.00060 -0.00250 0.01943 -0.00020 Pericarp thickness 0.00104 0.00241 -0.00030 0.00273 0.00529 0.00556 -0.00030 0.03820 Correlation with fruit yield 0.6549** 0.6645** 0.1538 0.9119** 0.1555 0.09296 0.0810 0.13789

Plant height No. of primary branches Fruits per truss Fruits per plant Average fruit weight Locules per fruit Fruit shape index Pericarp thickness

Diagonal values represent direct effects *Significant at 5% level of probability **Significant at 1% level of probability

Table 19. Path coefficient analysis of different characters towards fruit yield per plant in F3 population of tomato Character Plant height Number of primary branches 0.57054 0.05341 -0.04320 -0.02760 -0.01240 -0.00470 0.00128 -0.00170 Fruits per truss 0.00941 0.01340 0.05205 0.00421 -0.00010 0.04224 0.02908 0.03970 Fruits per plant -0.00410 0.58080 0.28950 0.90820 0.00600 0.00003 0.00126 0.00026 Average fruit weight 0.02756 0.00386 0.01154 0.03415 0.21556 -0.00280 0.00363 -0.00440 Locules per fruit 0.00026 -0.00420 -0.00600 -0.00110 0.03370 0.03881 -0.00800 0.01694 Fruit shape index 0.00354 0.00143 0.00043 0.00302 0.00283 -0.00210 0.02097 0.00039 Pericarp thickness 0.00088 -0.00210 -0.00070 -0.00430 0.00474 0.00554 0.00047 0.02576 Correlation with fruit yield 0.5585** 0.6541** 0.30422** 0.93837** 0.4368** 0.06428 0.0708 -0.08876

Plant height No. of primary branches Fruits per truss Fruits per plant Average fruit weight Locules per fruit Fruit shape index Pericarp thickness

0.00749 0.00646 0.00140 0.01978 -0.16610 -0.01280 0.03096 -0.15160

Diagonal values represent direct effects *Significant at 5% level of probability **Significant at 1% level of probability

The direct effect of primary branches on fruit yield was low in magnitude (0.048 and 0.059 for population M and population S respectively) but its correlation with fruit yield was high and significant to the tune of 0.74 and 0.71 in intra mating populations. Primary branches exerted high indirect effect through fruits per plant (0.573 and 0.648 for population M and S respectively) on fruit yield. Though the association of fruits per truss was high in magnitude of 0.795 and 0.530 for population M and population S respectively, its direct effect was low (0.1280 and 0.053 for population M and S respectively). The indirect effect of fruits per truss via fruits per plant on fruit yield was high (0.573 and 0.582 for population M and S respectively). Through plant height the indirect effect was negative where as the indirect effect through primary branches, average fruit weight, shape index was positive in intra mating populations. The association of fruits per plant with fruit yield was high, positive and significant in intra mating population (0.905 and 0.948 for population M and S respectively). The direct effect of the trait under consideration was also high (0.759 and 0.934 respectively for population M and S). Though the indirect effect via plant height was positive (0.0407) in population M, it was negative (-0.058) in population S. The locules per fruit, pericarp thickness, shape index and fruits per truss exhibited indirect effect on fruit yield in intra mating population. The direct effect of average fruit weight on fruit yield was of higher in magnitude (0.353 and 0.296 respectively for population M and S and its association with fruit yield was positively significant (0.406 and 0.494 for population M and S respectively). The indirect effect via other characters was negligible. The correlation between locules per fruit and fruit yield was positive (0.049 and 0.137 respectively for population M and S). Its direct effect an fruit yield was negative (-0.032) in population M but positive (0.027) in population S. The influence of locules per fruit viz pericarp thickness (-0.0677 and -0.0109 for population M and S respectively) and fruits per plant (0.00063 and ­0.0211 for population M and S respectively) was negative on fruit yield in intra mating population. The association between shape index and fruit yield was negative (-0.128) in population M but positive (0.164) in population S. Similarly the direct effect of it on fruit yield was negative (-0.0203) in population M, but positive (0.041) in population S. The indirect effect of shape index through other traits under study on fruit yield was very low in intra mating populations. It was observed that the pericarp thickness was positively and significantly associated with fruit yield (0.287) in population S, but the association turned to be non significant (0.127) in population M. The pericarp thickness had high magnitude of indirect effect on fruit yield through average fruit weight (0.1506 and 0.11 for population M and S respectively). The direct effect of it was negative (-0.130 and ­0.0316 for population M and S respectively) in both intra mating populations. 4.1.6.2 Inter mating populations The association between plant height and fruit yield (0.702 and 0.654 respectively for population M/S and S/M) was positive and significant and so also the direct effect of this trait on fruit yield was positive (0.045 and 0.004 for population M/S and S/M respectively). The indirect effect of plant height via primary branches was high (0.613 and 0.575 respectively for population M/S and S/M) where as its indirect effect viz., other characters under study was low in both the inter mating populations. The indirect effect of primary branches via fruits per plant on fruit yield was high and positive (0.483 and 0.545 respectively for population M/S and S/M) but its direct effect was low (0.031 and 0.039 for population M/S and S/M respectively) though it has positively and significantly correlated with fruit yield (0.56 and 0.66 respectively for population M/S and S/M). The magnitude of indirect effect of trait under study viz plant height (0.031 and 0.039 respectively for population M/S and S/M) and average fruit weight (0.028 and 0.0025 respectively for population M/S and S/M) followed that of fruits per plant. Fruits per truss exhibited low direct effect on fruit yield (0.037 and 0.023 respectively for population M/S and S/M) in both inter mating populations. It has positively and significantly associated with fruit yield (0.43) in population M/S but non-significant and positive associated

(0.153) in population S/M. Its indirect effect via fruits per plant (0.377 and 0.102 for population M/S and S/M respectively) on fruit yield was high whereas via other characters under study was low in the both inter mating populations. Fruits per plant exhibited high positive correlation with fruit yield (0.939 and 0.911 for population M/S and S/M respectively) and showed high direct effect on fruit yield (0.889 and 0.866 respectively for population M/S and S/M). The indirect effect via plant height (-0.016 and 0.009) was negative in both inter mating populations. Though average fruit weight was positively and significantly associated with fruit yield in population M/S (0.298), it was non-significant (0.155) in population S/M. The direct effect of this trait was high (0.228 and 0.259 in population M/S and S/M respectively) in both the inter mating populations. Locules per fruit showed positive correlation with fruit yield (0.10 and 0.09 respectively for population M/S and S/M). Its direct effect was 0.0059 for population M/S and 0.014 for population S/M. Its indirect effect was maximum through plant height (0.052) in population M/S and fruits per truss (0.051) in population S/M. The indirect effect of it through other characters under study was very meager. Though the shape index negatively correlated with fruit yield (-0.066) in population M/S, but it was positively correlated (0.081) in population S/M. Shape index had on direct effect of 0.016 and 0.019 for population M/S and S/M respectively. The indirect effect via other characters understudy was negligible Pericarp thickness exhibited positive correlation with fruits yield in inter mating populations (0.10 and 0.13 for population M/S and S/M respectively). Its direct effect was negative (-0.013) in population M/S though positive (0.038) in population S/M. Pericarp thickness had an indirect effect through average fruit weight (0.059 and 0.061 for population M/S and S/M respectively) in both the inter mating populations.

4.1.7 Selfed population (F3)

Even though, high and positively significant association (0.558) between plant height and fruit yield was observed, the direct effect of plant height on fruit yield was very low (0.0074). The indirect effect of plant height via primary branches (0.57) on fruit was high. The indirect effect of plant height via other characters viz., fruits per truss (0.0094) fruits per plant (-0.004), average fruit weight (0.027), locules per fruit (0.00026), shape index (0.0035) and pericarp thickness (0.00088) was very low on yield. The effect of primary branches via fruits per plant was high (0.580). The indirect effect of primary branches via locules per fruit (-0.0042) and pericarp thickness (-0.0021) was negative. The fruits per truss exhibited positive direct effect on fruits yield per plant (0.062) and it was also positively and significantly associated with fruit yield (0.304). The indirect effect of it via plant height (0.289) was higher than rest of the characters. The fruits per plant exhibited high direct effect (0.908) on yield per plant. The indirect effect via primary branches, (-0.0278) locules per fruit (-0.0011) and pericarp thickness (0.0043) was low negative and it was positive and low through rest of the characters. Average fruit weight showed positive and significant association with fruit yield (0.436) similarly it had showed positive direct effect (0.215) on average fruit weight. Except plant height which showed indirect effect of 0.166 on fruit yield, all other traits showed very negligible indirect effect on yield. The locules/fruit showed positive and direct effect (0.038) on the yield per plant. The indirect effect of locules per fruit viz other characters was very negligible. The character also associated positively with fruit yield. The association between the fruit shape index and yield per plant was very low and positive. The direct effect of the trait on yield per plant was 0.02097. Though the pericarp thickness was negatively associated with fruit yield (-0.00887) its direct effect on fruit yield was positive (0.0257).

4.1.8 Identification of superior segregants

The number of superior segregants in inter mating populations was higher (35) than in intra mating population (12). Out of 4 BIP populations, the population S/M showed higher number of superior segregants (21) followed by population M/S (14). In the range of 10501200 class interval, the higher number of superior segregants was observed (Table 20). Compared to BIPs the F3 showed lower number of superior segregants (8).

4.2 EXPERIMENT ­ II: DIALLEL MATING

The results obtained from the present investigation are presented under following headings. 4.2.1 Analysis of variance 4.2.2 Per se performance and magnitude of heterosis 4.2.3 Combining ability

4.2.1 Analysis of variance

The analysis of variance (Table 21) indicated significantly higher amount of variability among the genotypes for all the characters studied viz., plant height, number of primary branches, number of flowers per truss, number of fruits per truss, number of fruits per plant, average fruit weight, yield per plant, total soluble solids, number of locules per fruit, fruit shape index, pericarp thickness, ascorbic acid, total acidity, reducing sugar, non reducing sugar, total sugar, juice recovery and pulp content.

4.2.2 Per se performance and magnitude of heterosis

The pertinent data on per se performance of 28 entries and magnitude of heterosis are presented in Table 22 to 30. 4.2.2.1 Plant height (cm) The range for plant height was from 60.67 cm (CO-3) to 95.17 (Solan Vajra) in parents and from 51.57 cm (CO-3 x PKM-1) to 103.32 cm (Local x PKM-1) in F1s. The magnitude of heterosis for the trait under consideration ranged (Table 22) from ­ 26.33 (CO-3 x PKM-1) to 37.07 (L-15 x PKM-1), -35.00 (CO-3 x PKM-1) to 28.88 (Local x PKM-1) and ­29.69 (CO-3 x PKM-1) to 61.43 (L-15 x PKM-1) per cent over mid parent, better parent and commercial check respectively. Out of 21 hybrids, 12 exhibited significant heterosis over mid parent of which ten showed heterosis in the positive direction. Nine out of 21 hybrids, exhibited significant heterosis over the better parent, of which five showed heterosis in positive direction. Sixteen of the 21 hybrids showed significant positive heterosis over the commercial check and only one (CO-3 x PKM-1) showed significant negative heterosis. 4.2.2.2 Number of primary branches Among the parents, the number of primary branches ranged from 4.33 (PKM-1) to 6.32 (Local type). However for the hybrids, it was in the range of 3.45 (Solan Vajra x PKM-1) to 6.5 (S-22 x Local). The extent of heterosis exhibited by the F1s over their corresponding mid parent, better parent and commercial check ranged from ­34.6 (Solan Vajra x PKM-1) to 33.33 (L15xPKM-1), -44.50 (Solan Vajra x PKM-1) to 25.42 (L-15 x PKM-1) and ­5.73 (Solan Vajra x PKM-1) to 91.25 (Local type x PKM-1), respectively. Among the hybrids exhibiting significant heterosis over mid parent four showed significant positive heterosis. Thirteen hybrids showed significant heterosis over better parent of which only one hybrid showed significant hetero beltiosis in positive direction (Table 22). Out of 21 only fifteen hybrids showed significant heterosis over commercial check, and all of them showed positive economic heterosis.

Table 20. Frequency distribution of superior segregants for fruit yield in different segregating populations of tomato

Population

10501200

12001350

13501500

15001650

16501800

18001950

19502100

21002250

22502400

24002550

25502700

27002850

28503000

30003150

31503300

M

3

1

0

1

S

3

1

2

1

M/S

4

3

5

1

1

S/M

7

8

4

2

F3

5

-

2

1

Table 21. Analysis of variance in respect of 18 characters in a set of 7x7 half diallel crosses in tomato

4.2.2.3 Flowers per truss The average number of flowers per truss ranged from 3.27 (Sivap) to 5.42 (Local type) in parents and 3.27 (CO-3 x PKM-1) to 6.05 (Local x PKM-1) in crosses respectively (Table 23). Heterosis ranged between ­32.88 (CO-3 x PKM-1) to 33.5 (S-22 x L-15) per cent over mid parent, -39.52 (CO-3 x PKM-1) to 30.26 (S-22 x L-15) per cent over better parent and ­2.09 (CO-3 x PKM-1) to 63.17 (S2 x Local) per cent over commercial check. Out of 21 hybrids 14 over mid parent, 11 over better parent and 15 over commercial check showed significant heterosis. Further of the crosses showing significant heterosis over mid parent, better parent and commercial check 10, 4 and 15 respectively showed heterosis in the desired (positive) direction. 4.2.2.4 Fruits per truss The average fruits per truss ranged from 2.55 (Sivap) to 4.23 (Local type) for parents and 2.47 (S-22 x Sivap) to 4.99 (Local x PKM-1) for hybrids. The magnitude of heterosis for the trait under consideration ranged (Table 23) from ­ 16.87 (CO-3 x PKM-1) to 39.46 (Sivap x Local), -16.27 (L-15 x PKM-1) to 25.86 (L-15 x CO-3) and 2.62 (S-22 x CO-3, L-15 x Sivap) to 4.99 (Local x PKM-1) per cent over mid parent, better parent commercial check, respectively. Among hybrids showing significant heterosis nine, four and eight hybrids expressed significant positive heterosis over mid parent, better parent and standard check respectively. 4.2.2.5 Number of fruits per plant The results from per se performance of parents and hybrids revealed that, the parents had number of fruits per plant ranging from 32.67 (L-15) to 95 (Local type). However in the hybrids it was in the range of 25.00 (Sivap x Solan Vajra) and 125.33 (Local x PKM-1) (Table 24). For the trait under consideration, the heterosis observed was in the range of ­24.62 (Sivap x Solan Vajra) and 95.83 (Local x PKM-1) per cent over mid parent, -25.00 (Sivap x Solan Vajra) and 31.93 (Local x PKM­1) per cent over better parent and ­18.46 (Sivap x Solan Vajra) and 308.77 (Local x PKM-1) per cent over commercial check Out of 21 hybrids nine over mid parent, five over better parent and 13 over commercial check showed heterosis in the positive direction. 4.2.2.6 Average fruit weight (g) Average fruit weight ranged from 7.69 (Local type) to 34.27 (Solan Vajra) and from 6.71 (Sivap x Local type) and 37.6 (L-15 x PKM-1) in parents and hybrids respectively. The data on per cent heterosis (Table 24) revealed that out of 21 hybrids, 19, 17 and 17 hybrids showed significant heterosis over mid parent, better parent and commercial check respectively. Significant heterosis in the positive direction was observed in nine crosses over mid parent, five crosses over better parent and three over commercial check. 4.2.2.7 Locules per fruit Among the parents, the number of locules per fruit ranged from 2.55 (L-15) to 4.77 (S-22). However in the hybrids, it was in the range of 2.26 (SivapxLocal) to 4.59 (S-22xPKM1). The extent of per cent heterosis (Table 25) exhibited by the F1s over their corresponding mid parent, better parent and commercial check ranged from -31.66 (Sivap x PKM-1) to 32.4 (L-15 x Sivap), -47.63 (S-22 x Local) to 15.50 (L-15 x Sivap) and -27.33 (Sivap x Local) to 47.40 (S-22 x PKM-1), respectively. Among the hybrids exhibiting significant heterosis over mid parent, only two showed significant negative heterosis. Out of 21 hybrids only 2 showed significant negative heterosis over better parent. Only three hybrids showed significant economic heterosis and none showed negative heterosis over commercial check.

Table 22. Per se performance, nature and magnitude of heterosis for plant height and primary branches in tomato Sl. No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 Plant height (cm) Per se performance Per cent heterosis Female Male F1 MP BP CC 77.5 93.33 79.17 -7.32 -15.18** 8.00 77.5 60.67 84.67 22.56** 9.25 15.51* 77.5 89.73 89.07 6.52 -0.74 21.51** 77.5 95.17 94.23 9.15** -0.98 28.55** 77.5 80.17 93.75 18.92 16.94** 27.89** 77.5 79.33 79.50 1.38 0.21 8.45 93.3 60.67 90.08 16.99** -3.48 22.89** 93.3 89.73 93.67 2.33 0.36 27.51** 93.3 95.17 100.67 6.81 5.78 37.33** 93.3 80.17 102.33 17.96** 9.64 39.60** 93.3 79.33 118.33 37.07** 26.79** 61.43** 60.67 89.73 91.00 21.01** 1.41 25.12** 60.67 95.17 100.92 29.52* 6.04 37.85** 60.67 80.17 92.08 30.77** 14.86* 26.15** 60.67 79.33 51.57 -26.33** -35.00** -9.69** 89.73 95.17 78.07 -15.56** -17.97** 6.50 89.73 80.17 100.08 17.81** 11.53* 37.11** 89.73 79.33 90.00 6.47 0.30 20.30** 95.17 80.17 87.92 0.29 -7.62 19.94** 95.17 79.33 81.33 -6.78 -14.54** 10.95 80.17 79.33 103.32 29.55** 28.88** 40.60** 73.30 4.69 4.03 4.72 4.72 9.19 7.98 9.25 9.22 12.05 10.35 12.17 12.00 Number of primary branches Per se performance Per cent heterosis Female Male F1 MP BP CC 4.83 4.92 4.17 -14.53* -15.25* 13.93 4.83 4.67 4.07 -14.39* -15.86* 11.20 4.83 4.75 3.92 -18.26** -18.97** 7.10 4.83 6.22 4.67 -15.54** -24.93** 26.22** 4.83 6.32 6.50 16.59** 2.90 75.59** 4.83 4.33 4.85 5.82 0.34 32.50** 4.92 4.67 4.80 0.17 -2.37 31.14** 4.92 4.75 4.92 1.72 0.00 34.40** 4.92 6.22 4.70 -15.57** -24.40** 28.41** 4.92 6.32 5.88 -0.59 -11.61 60.65** 4.92 4.33 6.17 33.33** 25.42** 68.50** 4.67 4.75 4.75 0.88 0.00 29.08** 4.67 6.22 5.07 -6.89 -18.50** 38.52** 4.67 6.32 4.50 -18.06** -28.76** 25.11* 4.67 4.33 3.75 -16.67** -19.64** 2.45 4.75 6.22 4.58 -16.41** -26.27** 25.13** 4.75 6.32 6.17 11.45* -2.37 68.50** 4.75 4.23 4.25 -6.42 -10.53 14.24 6.22 6.32 5.23 -16.49** -17.15** 42.80** 6.22 4.33 3.45 -34.6** -44.50** -5.73 6.32 4.33 6.20 31.46** 10.82 69.30** 3.66 0.37 0.31 0.34 0.34 0.72 0.60 0.66 0.67 0.95 0.79 0.87 0.87

Crosses S-22 x L-15 S-22 x CO-3 S-22 x Sivap S-22 x Solan Vajra S-22 x Local S-22 x PKM-1 L-15 x CO-3 L-15 x Sivap L-15 x Solan Vajra L-15 x Local L-15 x PKM-1 CO-3 x Sivap CO-3 x Solan Vajra CO-3 x Local CO-3 x PKM-1 Sivap x Solan Vajra Sivap x Local Sivap x PKM-1 Solan Vajra x Local Solan Vajra x PKM-1 Local x PKM-1 MHTM-256(CC) S.Em± CD at 5% CD at 1%

*Significant at 5% level of probability **Significant at 1% level of probability MP ­ Mid parent, BP-Better parent, CC-Commercial check

Table 23. Per se performance, nature and magnitude of heterosis for flowers per truss and fruits per truss in tomato

Sl. No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21

Crosses S-22 x L-15 S-22 x CO-3 S-22 x Sivap S-22 x Solan Vajra S-22 x Local S-22 x PKM-1 L-15 x CO-3 L-15 x Sivap L-15 x Solan Vajra L-15 x Local L-15 x PKM-1 CO-3 x Sivap CO-3 x Solan Vajra CO-3 x Local CO-3 x PKM-1 Sivap x Solan Vajra Sivap x Local Sivap x PKM-1 Solan Vajra x Local Solan Vajra x PKM-1 Local x PKM-1 MHTM-256(CC)

S.Em± CD at 5% CD at 1%

Flowers per truss Per se performance Per cent heterosis Female Male F1 MP BP CC 3.50 3.33 4.56 33.5** 30.26** 36.52** 3.50 4.34 4.23 7.91 -2.46 26.64** 3.50 3.27 3.50 3.30** -0.01 4.79 3.50 3.33 4.00 17.03** 14.08* 17.64** 3.50 5.42 5.45 22.18** 0.55 63.17** 3.50 5.41 4.02 -9.69* -25.50** 20.35** 3.33 4.34 4.59 19.69** 5.84 37.4** 3.33 3.27 3.25 -1.51 -2.40 -2.69 3.33 3.33 4.21 26.33** 26.45** 26.04** 3.33 5.42 4.33 -1.18 -20.22** 29.6** 3.33 5.41 4.06 -7.09 -24.91** 21.55** 4.43 3.27 4.31 13.36* -0.54 29.04** 4.43 3.33 3.29 -14.05** -24.06** -1.49 4.43 5.42 5.59 14.48** 3.01 67.3** 4.43 5.41 3.27 -32.88** -39.52** -2.09 3.27 3.33 3.39 2.63 1.80 1.49 3.27 5.42 5.39 23.96** -0.61 61.3** 3.27 5.41 4.05 -6.68 -25.09** 21.2** 3.33 5.42 5.35 22.36** -1.29 60.17** 3.33 5.41 3.42 -21.38** -36.74** 2.39 5.42 5.41 6.05 11.67** 11.49** 81.1** 3.34

0.22 0.43 0.56 0.19 0.37 0.49 0.21 0.41 0.54 0.21 0.41 0.54

Fruits per truss Per se performance Per cent heterosis Female Male F1 MP BP CC 2.77 2.60 3.17 17.87* 14.32* 16.11* 2.77 2.78 2.62 -5.64 -5.87 -4.02 2.77 2.55 2.47 -7.02 -10.71 -9.52 2.77 2.60 2.65 -1.30 -4.33 -2.93 2.77 4.23 4.34 23.65** 1.92 58.97** 2.77 3.40 2.92 -5.35 -14.12* 6.95 2.60 2.78 3.28 21.66** 25.86** 20.14* 2.60 2.55 2.62 1.55 0.51 -4.02 2.60 2.60 3.10 19.03* 19.10** 13.85 2.60 4.23 4.13 20.92** -2.36 51.28** 2.60 3.40 2.85 -5.16 -16.27* 4.39 2.78 2.55 2.87 7.63 3.11 5.12 2.78 2.60 2.77 2.79 -0.60 1.46 2.78 4.23 4.53 29.22** 7.09 65.39** 2.78 3.40 2.57 -16.87** -24.41** -5.86 2.55 2.60 2.47 -3.95 -4.87 -9.52 2.55 4.23 4.73 39.46** 11.73 73.20** 2.55 3.40 2.92 -1.85 -14.12* 6.95 2.60 4.23 4.74 38.63** 11.89 73.62** 2.60 3.40 2.97 -1.00 -1.29 6.22 4.23 3.40 4.99 30.83** 17.95** 82.78** 2.73

0.24 0.47 0.61 0.20 0.39 0.51 0.22 0.43 0.56 0.22 0.43 0.56

*Significant at 5% level of probability **Significant at 1% level of probability MP ­ Mid parent, BP-Better parent, CC-Commercial check

Table 24. Per se performance, nature and magnitude of heterosis for fruits per plant and average fruit weight in tomato

Sl. No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21

Crosses S-22 x L-15 S-22 x CO-3 S-22 x Sivap S-22 x Solan Vajra S-22 x Local S-22 x PKM-1 L-15 x CO-3 L-15 x Sivap L-15 x Solan Vajra L-15 x Local L-15 x PKM-1 CO-3 x Sivap CO-3 x Solan Vajra CO-3 x Local CO-3 x PKM-1 Sivap x Solan Vajra Sivap x Local Sivap x PKM-1 Solan Vajra x Local Solan Vajra x PKM-1 Local x PKM-1 MHTM-256(CC)

S.Em± CD at 5% CD at 1%

Fruits per plant Per se performance Per cent heterosis MP BP CC Female Male F1 34.67 32.67 41.33 22.77** 19.23** 34.78** 34.67 28.67 29.00 -8.47 -16.35* -5.41 34.67 33.33 40.67 19.61** 17.31* 32.64** 34.67 33.00 30.33 -10.34 -12.50 -1.07 34.67 95.00 121.33 87.15** 27.72** 295.72** 34.67 33.00 30.67 -9.36 -11.54 0.032 32.67 28.67 39.33 28.26** 20.41** 28.2** 32.67 33.33 35.67 8.08 7.00 16.30** 32.67 33.00 27.33 -16.75* -17.17* -10.86** 32.67 95.00 101.33 58.75** 6.67 230.49** 32.67 33.00 26.67 -18.78** -19.79** -13.01** 28.67 33.33 39.33 26.55** 18.00** 28.27** 28.67 33.00 34.67 12.43 5.05 13.07** 28.67 95.00 75.00 -21.29** -21.05** 144.61** 28.67 33.00 28.67 -7.03 -13.13 -6.5** 33.33 33.00 25.00 -24.62** -25.00** -18.46** 33.33 95.00 103.00 60.52** 8.42 235.9** 33.33 33.00 33.33 0.50 0.00 8.70** 33.33 95.00 76.67 19.79** -19.03** 150.06** 33.33 33.00 28.33 -14.14* -14.14* -7.59** 95.00 33.00 125.33 95.83** 31.93** 308.77** 30.66

2.35 4.60 6.03 2.03 3.98 5.21 2.32 4.54 5.96 2.32 4.54 5.96

Average fruit weight (g) Per se performance Per cent heterosis Female Male F1 MP BP CC 29.33 23.20 32.42 23.41** 10.51** 1.94 29.33 21.72 29.92 17.1** 2.00 -5.91** 29.33 29.47 31.73 7.94** 7.69** -2.20 29.33 34.27 28.55 -10.22** -16.68** -10.22** 29.33 7.69 7.86 -57.55** -73.22** -75.28** 29.33 27.30 32.40 14.42** 18.68** 1.88 23.20 21.77 26.83 19.35** 15.66** -15.62** 23.20 29.47 25.97 -1.39 -11.88** -18.33** 23.20 34.27 25.87 -9.98** -24.57** -1.86 23.20 7.69 8.57 -44.53** -63.07** -73.05** 23.20 27.30 37.60 48.91** 37.73** 18.26** 21.7 29.47 26.73 4.36 -9.28** -15.89** 21.7 34.27 34.07 21.59** -0.56 7.13** 21.7 7.69 8.81 -40.15** -59.51** -72.29** 21.7 27.30 22.33 -8.97** -18.19** -29.77** 29.47 34.27 35.30 10.77** 3.02 11.00** 29.47 7.69 6.71 -63.87** -77.22** -78.89** 29.47 27.30 30.13 6.17** 2.26 -4.30* 34.27 7.69 8.88 -57.65** -74.08** -71.80** 34.27 27.30 28.20 -8.39** -17.70** -11.32** 7.69 27.30 7.43 -57.51** -72.77** -75.95** 31.80

0.66 1.29 1.69 0.59 1.15 1.51 0.71 1.39 1.83 0.71 1.38 1.84

*Significant at 5% level of probability **Significant at 1% level of probability MP ­ Mid parent, BP-Better parent, CC-Commercial check

Table 25. Per se performance, nature and magnitude of heterosis for locules per fruit and fruit shape index in tomato

Sl. No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21

Crosses S-22 x L-15 S-22 x CO-3 S-22 x Sivap S-22 x Solan Vajra S-22 x Local S-22 x PKM-1 L-15 x CO-3 L-15 x Sivap L-15 x Solan Vajra L-15 x Local L-15 x PKM-1 CO-3 x Sivap CO-3 x Solan Vajra CO-3 x Local CO-3 x PKM-1 Sivap x Solan Vajra Sivap x Local Sivap x PKM-1 Solan Vajra x Local Solan Vajra x PKM-1 Local x PKM-1 MHTM-256(CC)

S.Em± CD at 5% CD at 1%

Locules per fruit Per se performance Per cent heterosis Female Male F1 MP BP CC 4.77 2.55 3.61 -1.37 -24.37* 16.07 4.77 3.50 4.55 9.91 -4.75 46.22* 4.77 3.42 4.37 6.59 -8.25 40.53* 4.77 3.30 3.48 -13.87 -27.16* 11.89 4.77 2.70 2.50 -33.1* -47.63** -19.64 4.77 3.76 4.59 7.54 -3.91 47.40* 2.55 3.50 2.84 -6.06 -18.86 -8.67 2.55 3.42 3.95 32.40* 15.50 27.00 2.55 3.30 2.37 -18.93 -28.18* -23.79 2.55 2.70 2.39 -8.77 -11.36 -23.15 2.55 3.76 3.27 3.86 -12.86 5.14 3.50 3.42 3.27 -5.59 -6.67 5.14 3.50 3.33 3.39 -0.29 -3.14 9.00 3.50 2.70 2.27 -26.88 -35.24* -27.00 3.50 3.76 3.63 0.14 -3.28 16.72 3.42 3.30 3.62 7.74 5.85 16.70 3.42 2.70 2.26 -26.25 -34.02* -27.33 3.42 3.76 2.47 -31.16* -34.54* -20.50 3.30 2.70 2.43 -18.89 -26.26* -21.80 3.30 3.76 3.55 0.52 -5.59 14.14 2.70 3.76 2.67 -17.29 -28.93* -14.14 3.11

0.54 1.05 1.39 0.45 0.88 1.15 0.48 0.94 2.41 0.48 0.95 2.42

Fruit shape index (L/D) Per se performance Per cent heterosis Female Male F1 MP BP CC 0.77 0.95 0.86 -0.19 -9.79** -5.49 0.77 0.91 0.84 -0.59 -8.39** -7.69* 0.77 0.74 0.76 0.44 -1.73 -16.48** 0.77 0.88 0.77 -6.26 -12.12** -14.12** 0.77 0.88 0.84 1.61 -4.91 -7.69* 0.77 0.77 0.73 -4.76 -4.76 -19.78** 0.95 0.91 0.91 -2.14 -4.20 0.00 0.95 0.74 0.82 -3.35 -14.34** -9.51** 0.95 0.88 0.83 -9.82* -13.29** -8.34** 0.95 0.88 0.94 2.36 -1.40 3.02 0.95 0.77 0.73 -15.28** -23.43** -19.78** 0.91 0.74 0.83 0.61 -9.12** -8.34** 0.91 0.88 0.89 -1.12 -2.92 -2.01 0.91 0.88 0.99 10.2** 8.93** 8.79** 0.91 0.77 0.73 -13.66** -20.4** -19.78** 0.74 0.88 0.71 -11.75** -18.94** -21.97** 0.74 0.88 1.02 25.93** 15.47** 12.08** 0.74 0.77 0.72 -3.98 -6.06* -20.88** 0.88 0.88 0.87 1.32 -1.14 -6.56 0.88 0.77 0.83 0.2 -6.06* -8.34** 0.88 0.77 0.85 2.42 -4.15 -6.59* 0.91

0.032 0.06 0.08 0.031 0.05 0.08 0.03 0.06 0.08 0.03 0.06 0.08

*Significant at 5% level of probability **Significant at 1% level of probability MP ­ Mid parent, BP-Better parent, CC-Commercial check

4.2.2.8 Fruit shape index The results from per se performance of parents and hybrids (Table 25) revealed that, the parent values ranged from 0.74 (Sivap) to 0.95 (L-15). However, in the hybrids it was in the range of 0.71 (Sivap x Solan Vajra) to 1.02 (Sivap x Local). The magnitude of heterosis for the trait under consideration ranged from -15.28 (L-15 x PKM-1) to 25.93 (Sivap x Local), -23.43 (L-15 x PKM-1) to 15.47 (Sivap x Local) and -20.88 (Sivap x PKM-1) to 12.08 (Sivap x Local) per cent over mid parent, better parent and commercial check, respectively. Among the 21 hybrids, 6, 12 and 17 hybrids showed significant heterosis over mid parent, better and commercial check, respectively. Only two each showed positive heterosis over mid parent, better parent and commercial check respectively. 4.2.2.9 Pericarp thickness (mm) Among the parents, the pericarp thickness ranged from 2.6 (Local type) to 4.2 (CO3). However for the hybrids it was in the range of 2.2 (CO-3 x Local) to 4.8 (S-22 x L-15) (Table 26). Out of 21 hybrids under study 12, 11 and 13 showed significant heterosis over mid parent, better parent and commercial check respectively and among them five, two and seven hybrids recorded positive heterosis over mid parent, better parent and commercial check respectively. 4.2.2.10 Ascorbic acid (mg/100 g) The ascorbic acid content among parents ranged from 16.42 (PKM-1) to 27.00 (Local type) and among F1s it was between 17.08 (S-22 x L-15) and 3.92 (L-15 x Local) (Table 26). Heterosis ranged from -33.08 (S-22 x Solan Vajra) to 46.03 (Local x PKM-1) per cent over mid parent, -35.84 (S-22 x Solan Vajra) to 58.13 (Sivap x Local) per cent over better parent and 14.50 (S-22 x Solan Vajra) to 31.70 (Local x PKM-1) per cent over commercial check. Of all the hybrids under study, 16 of them over mid parent and 13 over better parent and all of them over commercial check showed significant heterosis. Further, out of the total crosses showing significant heterosis over mid parent, better parent and commercial check, 13, 8 and all of them respectively showed positive heterosis. 4.2.2.11 Total acidity (%) The results from per se performance of parents and hybrids (Table 27) revealed that, the parents ranged from 0.27 (S-22) to 0.43 (CO-3) while hybrids from 0.27 (S-22 x Solan Vajra and CO-3 x PKM-1) to 0.44 (S-22 x PKM-1) with respect to total acidity per cent in fruits. The data on per cent heterosis revealed that, more than fifty per cent of hybrids showed significant heterosis over mid parent, better parent and commercial check. Significant heterosis in positive direction was observed in eight crosses over mid parent, five crosses over better parent and 19 crosses over commercial check. 4.2.2.12 Reducing sugar (%) The reducing sugar ranged from 2.23 (CO-3) to 2.82 (S-22) among parents and 2.14 (L-15 x CO-3) to 2.95 (S-22 x L-15) among F1s (Table 27). The magnitude of heterosis for the trait under consideration ranged from -19.80 (L-15 x Local) to 28.21 (CO-3 x PKM-1), -23.30 to L-15 x Local) to 27.85 (CO-3 x PKM-1) and -13.7 (L-15 x Local) to 18.95 (S-22 x L-15) per cent over mid parent, better parent and commercial check respectively. Nineteen hybrids exhibited significant heterosis over mid parent of which seven showed heterosis in positive direction. Seventeen out of 21 hybrids showed significant heterosis of which three recorded positive heterosis over better parent. Fifteen hybrids showed significant heterosis of which six exhibited positive heterosis over standard check.

Table 26. Per se performance, nature and magnitude of heterosis for pericarp thickness and ascorbic acid in tomato

Sl. No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21

Crosses S-22 x L-15 S-22 x CO-3 S-22 x Sivap S-22 x Solan Vajra S-22 x Local S-22 x PKM-1 L-15 x CO-3 L-15 x Sivap L-15 x Solan Vajra L-15 x Local L-15 x PKM-1 CO-3 x Sivap CO-3 x Solan Vajra CO-3 x Local CO-3 x PKM-1 Sivap x Solan Vajra Sivap x Local Sivap x PKM-1 Solan Vajra x Local Solan Vajra x PKM-1 Local x PKM-1 MHTM-256(CC)

S.Em± CD at 5% CD at 1%

Pericarp thickness (mm) Per se performance Per cent heterosis Female Male F1 MP BP CC 3.4 3.9 4.8 31.5** 23.7** 41.17** 3.4 4.2 3.8 0.00 -9.52 11.76 3.4 3.6 3.8 8.57 5.55 11.76 3.4 4.4 4.08 4.61 -7.29 20.00** 3.4 2.4 2.6 -17.3** -23.52** -23.52** 3.4 3.6 4.0 14.2* 11.11 17.64** 3.9 4.2 3.8 -6.17 -9.52 11.76 3.9 3.6 3.4 -9.33 -12.82* 0.00 3.9 4.4 3.68 -11.32** -16.36** 8.23 3.9 2.4 2.2 -30.15** -43.5** -35.29** 3.9 3.6 4.4 17.33** 12.82* 29.41** 4.2 3.6 4.4 12.82** 4.76 29.41** 4.2 4.4 4.6 6.97 4.54 35.29** 4.2 2.4 2.2 -33.33** -47.6** -35.29** 4.2 3.6 3.6 -7.69 -14.28** 5.88 3.6 4.4 4.42 10.5* 0.45 30.00** 3.6 2.4 2.4 -20.0** -33.33** -29.41** 3.6 3.6 3.8 5.55 5.55 11.76 4.4 2.4 2.4 -29.41** -45.45** -29.41** 4.4 3.6 3.6 -10.00* -9.18 5.88 2.4 3.6 2.2 -26.66** -38.88** -35.29** 3.4

0.19 0.37 0.48 0.17 0.33 0.43 0.21 0.41 0.54 0.21 0.42 0.55

Ascorbic acid (mg/100 g) Per se performance Per cent heterosis Female Male F1 MP BP CC 20.73 21.05 17.08 -18.29** -18.97** 17.08** 20.73 25.58 17.25 -25.51** -32.57** 17.25** 20.73 16.92 21.83 15.98** 5.31 21.83** 20.73 22.60 14.50 -33.08** -35.84** 14.50** 20.73 27.00 24.75 3.70 -8.33* 24.75** 20.73 16.42 22.17 19.34** 6.91 22.17** 21.08 25.58 22.60 -3.14 -2.20 22.60** 21.08 16.92 25.58 34.65** 21.34** 25.58** 21.08 22.60 21.25 -2.71 -1.78 21.25** 21.08 27.00 30.92 28.6** 20.62** 30.92** 21.08 16.42 21.25 13.33** 0.79 21.25** 25.58 16.92 30.83 45.1** 20.59** 30.83** 25.58 22.60 31.42 30.4** 22.80** 31.42** 25.58 27.00 26.17 -0.48 2.28 26.17** 25.58 16.42 22.33 6.35 12.00** 22.33** 16.92 22.60 26.00 31.59** 18.72** 26.00** 16.92 27.00 26.75 21.82** 58.13** 26.75** 16.92 16.42 20.67 24.00** 25.89** 20.67** 22.60 27.00 26.92 8.53* 6.35 26.92** 22.60 16.42 24.50 25.59** 8.41 24.50** 27.00 16.42 31.70 46.03** 17.41** 31.70** 21.20

1.04 2.04 2.67 0.91 1.78 2.33 1.03 2.01 2.64 1.03 2.01 2.62

*Significant at 5% level of probability **Significant at 1% level of probability MP ­ Mid parent, BP-Better parent, CC-Commercial check

Table 27. Per se performance, nature and magnitude of heterosis for total acidity and reducing sugar in tomato

Sl. No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21

Crosses S-22 x L-15 S-22 x CO-3 S-22 x Sivap S-22 x Solan Vajra S-22 x Local S-22 x PKM-1 L-15 x CO-3 L-15 x Sivap L-15 x Solan Vajra L-15 x Local L-15 x PKM-1 CO-3 x Sivap CO-3 x Solan Vajra CO-3 x Local CO-3 x PKM-1 Sivap x Solan Vajra Sivap x Local Sivap x PKM-1 Solan Vajra x Local Solan Vajra x PKM-1 Local x PKM-1 MHTM-256(CC)

S.Em± CD at 5% CD at 1%

Total acidity (%) Per se performance Per cent heterosis Female Male F1 MP BP CC 0.27 0.36 0.39 24.87** 10.25** 50.00** 0.27 0.43 0.34 -2.83 -20.77** 30.7** 0.27 0.34 0.35 15.22** 3.92 34.6** 0.27 0.40 0.27 -20.4** -32.77** 3.84 0.27 0.35 0.34 8.02** -3.81 30.7** 0.27 0.37 0.44 36.46** 10.00** 69.2** 0.36 0.43 0.39 -1.27 -10.00** 50.00** 0.36 0.34 0.38 8.13** 5.61* 46.1** 0.36 0.40 0.43 14.16** 8.40** 65.3** 0.36 0.35 0.34 -4.72 -5.61* 30.7** 0.36 0.37 0.38 4.15 2.73 46.1** 0.43 0.34 0.42 9.46** -2.31 61.5** 0.43 0.40 0.27 -34.94** -37.69** 3.84 0.43 0.35 0.34 -14.04 -22.31** 30.7** 0.43 0.37 0.27 -33.6** -38.46** 3.84 0.34 0.40 0.34 -18.60** -15.13** 30.7** 0.34 0.35 0.39 12.08** 10.48** 50.00** 0.34 0.37 0.34 -3.80 -7.27** 30.7** 0.4 0.35 0.38 2.80 -3.36* 46.1** 0.4 0.37 0.33 -12.66** -15.97** 26.9** 0.35 0.37 0.37 4.19 1.82 42.3** 0.26

0.013 0.025 0.033 0.01 0.02 0.026 0.01 0.02 0.026 0.01 0.02 0.026

Reducing sugar (%) Per se performance Per cent heterosis Female Male F1 MP BP CC 2.82 2.79 2.95 5.05** 4.49* 18.95** 2.82 2.23 2.29 -9.11** -18.68** -7.66** 2.82 2.54 2.34 -12.56** -16.90** -5.64* 2.82 2.64 2.53 -7.45** -10.40** 2.01 2.82 2.55 2.82 5.09** 0.00 13.7** 2.82 2.24 2.72 7.51** -3.55 9.67** 2.79 2.23 2.48 -1.00 -10.99** 0.00 2.79 2.34 2.26 -15.22** -19.00** -8.87** 2.79 2.64 2.34 -13.81** -16.13** -5.64* 2.79 2.55 2.14 -19.80** -23.30** -13.7* 2.79 2.24 2.34 -6.83** -16.01** -5.64* 2.23 2.54 2.24 -6.01** -11.81** -9.67** 2.23 2.64 2.48 1.92 -6.06** 0.00 2.23 2.55 2.80 17.32** 9.95** 12.9** 2.23 2.24 2.86 28.21** 27.85** 15.32** 2.54 2.64 2.25 -12.74** -14.83** -8.87** 2.54 2.55 2.42 -4.85* -4.92** -2.42 2.54 2.24 2.48 3.77* -2.36 0.00 2.64 2.55 2.28 -12.08** -13.61** -8.1** 2.64 2.24 2.22 -9.02** -15.91** -10.5** 2.55 2.24 2.57 7.24** 0.79 3.63** 2.48

0.06 0.11 0.15 0.05 0.09 0.13 0.06 0.12 0.5 0.06 0.12 0.15

*Significant at 5% level of probability **Significant at 1% level of probability MP ­ Mid parent, BP-Better parent, CC-Commercial check

4.2.2.13 Non-reducing sugar (%) The non-reducing sugar content among parents ranged from 0.16 (S-22) to 0.57 (CO3), among F1s, it was from 0.17 (S-22 x Solan Vajra and CO-3 x PKM-1) to 0.60 (CO-3 x Sivap). The heterosis ranged from -68.12 (CO-3 x PKM-1) to 105.99 (Sivap x Solan Vajra), 70.35 (CO-3 x PKM-1) to 107.23 (Sivap x Solan Vajra) and -61.3 (S-22 x Solan Vajra and CO-3 x PKM-1) to 36.36 (CO-3 x Sivap) per cent over mid parent, etter parent and commercial check respectively. Of the total number of hybrids showing significant heterosis nine, four and two hybrids showed positive heterosis over mid parent, better parent and commercial check respectively (Table 28). 4.2.2.14 Total sugar (%) The total sugar content among the parents ranged from 2.75 (PKM-1) to 3.00 (L-15) and among F1s, it was from 2.64 (L-15 x Local) to 3.66 (S-22 x L-15) (Table 28). More than 50 per cent of hybrids under investigation showed significant heterosis over mid parent, better parent and commercial check. The heterosis ranged between -9.06 (L15 x Local) and 8.72 (CO-3 x PKM-1), -11.84 (L-15 x Local) and 8.02 (CO-3 x Local) and 11.2 (Solan Vajra x PKM-1) and 9.67 (S-22 x CO-3) per cent respectively over mid parent, better parent and commercial check. Out of total hybrids showing significant heterosis, eight and five showed positive heterosis over mid parent and better parent respectively and six showed positive heterosis over commercial check. 4.2.2.15 Juice recovery percentage (%) The results from per se performance of parents and hybrids (Table 29) revealed that, the parents had a range from 27.92 (CO-3) to 39.5 (PKM-1), whereas F1s ranged from 19.37 (Local x PKM-1) to 40.67 (S-22 x CO-3). For the trait under consideration the heterosis was observed in the range of -43.78 (Local x PKM-1) to 23.05 (S-22 x CO-3) per cent over mid parent, -50.97 (Local x PKM-1) to 14.41 (L-15 x Local) per cent over better parent and -14.66 (Local x PKM-1) to 79.16 (S-22 x CO-3) per cent over commercial check. Out of 21 hybrids, 13 and 15 hybrids showed significant heterosis over mid parent and commercial check and 12 showed significant heterosis over better parent. Of the hybrids showing significant heterosis, five, one and 15 recorded positive heterosis. 4.2.2.16 Pulp content (%) Average pulp content ranged from 58.1 (S-22) to 67.00 (L-15) among parents and 37.89 (S-22 x Local) to 76.61 (Local x PKM-1) among F1s (Table 29). Except seven all the hybrids under study showed significant heterosis over mid parent. Except nine and five hybrids, rest recorded heterosis over better parent and commercial check respectively. The heterosis percentage ranged from -7.11 (S-22 x Local) to 22.69 (Local x PKM-1) over mid parent, -12.97 (S-22 x Local) to 19.54 (S-22 x PKM-1) over better parent and -23.56 (S-22 x Local) to -5.24 (S-22 x L-15) over commercial check. Out of total number of hybrids showing significant heterosis 13 and 10 hybrids respectively showed positive heterosis over mid parent and better parent, however none hybrid exhibited positive heterosis over commercial check. 4.2.2.17 Total soluble solids (%) The total soluble solids ranged from 3.47 (S-22) to 6.23 (Local type) and 4.00 (S-22 x Sivap) to 6.37 (Local x PKM-1) in parents and F1s respectively. The data on per cent heterosis (Table 30) revealed that majority of hybrids had significant heterosis over mid parent and commercial check. Significant heterosis in positive direction was observed in 11 crosses over mid parent, in four crosses over better parent and in nine crosses over the commercial check.

Table 28. Per se performance, nature and magnitude of heterosis for non-reducing sugar and total sugar in tomato

Sl. No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21

Crosses S-22 x L-15 S-22 x CO-3 S-22 x Sivap S-22 x Solan Vajra S-22 x Local S-22 x PKM-1 L-15 x CO-3 L-15 x Sivap L-15 x Solan Vajra L-15 x Local L-15 x PKM-1 CO-3 x Sivap CO-3 x Solan Vajra CO-3 x Local CO-3 x PKM-1 Sivap x Solan Vajra Sivap x Local Sivap x PKM-1 Solan Vajra x Local Solan Vajra x PKM-1 Local x PKM-1 MHTM-256(CC)

S.Em± CD at 5% CD at 1%

Non-reducing sugar (%) Per se performance Per cent heterosis Female Male F1 MP BP CC 0.16 0.21 0.19 2.70 -9.52 -56.8** 0.16 0.57 0.42 14.56 -16.74 -4.54 0.16 0.28 0.34 55.73** 22.89 -22.72* 0.16 0.28 0.17 -21.21 -38.10** -61.3** 0.16 0.33 0.23 -5.41 -30.00** -47.72** 0.16 0.44 0.24 -25.51 -50.68** -45.45** 0.21 0.57 0.35 -9.79 -38.37** -20.45 0.21 0.28 0.41 68.49** 48.19** -6.81 0.21 0.28 0.49 98.64** 73.81** 11.36 0.21 0.33 0.49 80.37** 47.00** 11.36 0.21 0.49 0.45 27.01** -9.46 2.72 0.57 0.28 0.60 40.39** 4.07 36.36** 0.57 0.28 0.46 8.59 -19.19 4.54 0.57 0.33 0.24 -46.32** -57.56** -45.45** 0.57 0.49 0.17 -68.12** -70.35** -61.30** 0.28 0.28 0.57 105.99** 107.23** 29.54** 0.28 0.33 0.40 30.05** 19.00 -9.09 0.28 0.49 0.37 -4.76** -25.68** -15.9 0.28 0.33 0.39 27.17* 17.00 -11.36 0.28 0.49 0.37 -3.45 -24.32** -15.90 0.33 0.49 0.42 0.81 -15.54 -4.45 0.44

0.047 0.09 0.12 0.04 0.08 0.10 0.05 0.10 0.13 0.05 0.10 0.13

Total sugar (%) Per se performance Per cent heterosis Female Male F1 MP BP CC 2.98 3.00 3.16 5.69** 5.33** 7.48** 2.98 2.83 2.72 -6.31** -8.77** 9.67** 2.98 2.81 2.71 -6.44** -9.06** -7.82** 2.98 2.93 2.71 -8.46** -9.17** -7.82** 2.98 2.81 3.04 4.95** 2.01 3.4* 2.98 2.75 3.01 5.12** 1.12 2.0 3.00 2.83 2.85 -2.06* -4.89** -3.06* 3.00 2.81 2.64 -7.34** -10.22** -8.50** 3.00 2.93 2.86 -3.48** -4.56** -2.72** 3.00 2.81 2.64 -9.06** -11.89** -10.2** 3.00 2.75 2.82 -1.85 -5.89** -4.08** 2.83 2.81 2.87 1.77 1.53 -2.58** 2.83 2.93 2.97 3.24** 1.36 1.02* 2.83 2.81 3.05 8.27** 8.02** 3.74** 2.83 2.75 3.03 8.72** 7.31** 3.06* 2.81 2.93 2.87 -0.23 -2.27 -2.38 2.81 2.81 2.83 0.47 0.47 -3.74** 2.81 2.75 2.88 3.47** 2.37 -2.04 2.93 2.81 2.69 -6.26** -8.18** -8.50** 2.93 2.75 2.61 -8.09** -10.91** -11.22** 2.81 2.75 2.97 6.83** 5.64** 1.02 2.94

0.03 0.058 0.077 0.03 0.06 0.08 0.04 0.08 0.10 0.04 0.08 0.11

*Significant at 5% level of probability **Significant at 1% level of probability MP ­ Mid parent, BP-Better parent, CC-Commercial check

Table 29. Per se performance, nature and magnitude of heterosis for juice recovery percentage and pulp content in tomato

Sl. No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21

Crosses S-22 x L-15 S-22 x CO-3 S-22 x Sivap S-22 x Solan Vajra S-22 x Local S-22 x PKM-1 L-15 x CO-3 L-15 x Sivap L-15 x Solan Vajra L-15 x Local L-15 x PKM-1 CO-3 x Sivap CO-3 x Solan Vajra CO-3 x Local CO-3 x PKM-1 Sivap x Solan Vajra Sivap x Local Sivap x PKM-1 Solan Vajra x Local Solan Vajra x PKM-1 Local x PKM-1 MHTM-256(CC)

S.Em± CD at 5% CD at 1%

Juice recovery (%) Per se performance Per cent heterosis Female Male F1 MP BP CC 38.18 30.3 25.43 -25.72** -33.38** 12.02 38.18 27.92 40.67 23.05** 6.52 79.18** 38.18 34.4 34.77 -4.99 -8.93 53.71** 38.18 34.98 39.83 8.9 4.34 75.45** 38.18 29.39 37.80 11.88* -0.99 66.50** 38.18 39.5 33.97 -12.54* -14.01* 49.17** 30.3 27.92 27.07 -7.02 -10.67 19.25* 30.3 34.4 21.40 -33.85** -37.79** -5.72 30.3 34.98 34.67 6.21 -0.89 52.73** 30.3 29.39 34.67 16.15** 14.41* 52.73** 30.3 39.5 29.43 -15.66* -25.44** 29.64** 27.92 34.4 34.70 11.35* 0.87 52.86** 27.92 34.98 28.30 -10.02 -19.09** 24.66** 27.92 29.39 25.40 -11.37* -13.59* 4.92 27.92 39.5 36.33 7.78 -8.02 60.06** 34.4 34.98 33.63 -3.04 -3.84 48.14** 34.4 29.39 23.90 25.07** -30.52** 5.28 34.4 39.5 35.60 -3.65 -9.87* 56.82** 34.98 29.39 20.67 -35.7** -40.91** -8.94 34.98 39.5 27.47 -26.24** -30.46** 21.01* 29.39 39.5 19.37 -43.78** -50.97** -14.66 22.70

2.18 4.27 5.60 1.83 3.58 4.70 1.99 3.90 5.11 1.99 3.91 5.13

Pulp content (%)

Per se performance Female Male F1 58.10 67.00 71.73 58.10 66.87 62.77 58.10 61.23 63.67 58.10 63.57 62.23 58.10 66.48 57.86 58.10 58.40 69.81 67.00 66.87 70.62 67.00 61.23 76.17 67.00 63.57 63.15 67.00 66.48 75.03 67.00 58.40 68.73 66.87 61.23 64.43 66.87 63.57 70.15 66.87 66.48 71.45 66.87 58.40 61.96 61.23 63.57 63.23 61.23 66.48 74.58 61.23 58.40 63.03 63.57 66.48 76.35 63.57 58.40 69.73 66.48 58.40 76.61 75.70

2.17 4.25 5.57

Per cent heterosis MP BP CC 14.68** 7.06* -5.24 0.45 -6.13* -17.1** 6.70* 3.97 -15.84** 2.30 -2.10 -17.79** -7.11* -12.97** -23.56** 19.85** 19.54** -7.65** 5.51 5.40 -6.58* 18.79** 13.68** 0.55 -3.27 -5.75 -16.57** 12.43** 11.99** -0.88 19.62** 2.59 -9.2** 0.60 -3.64 -14.88** 7.56* 4.90 -7.33** 7.11* 6.80* -5.65* -1.08 37.34** -18.1** 1.33 -0.53 -16.47** 16.80** 12.19** -14.79** 15.37** 2.93 -16.73** 17.42** 14.85** 0.86 14.35** 9.70** -7.88** 22.69** 15.23** 1.32

1.86 3.64 4.78 2.05 4.01 5.27 2.05 4.01 5.26

*Significant at 5% level of probability **Significant at 1% level of probability MP ­ Mid parent, BP-Better parent, CC-Commercial check

Table 30. Per se performance, nature and magnitude of heterosis for fruit yield and total soluble solids in tomato Sl. No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 Total soluble solids (°Brix) Per se performance Per cent heterosis Female Male F1 MP BP CC 3.47 4.30 4.50 15.78** 4.65 -1.3 3.47 4.20 4.55 18.78** 8.50 -0.22 3.47 4.48 4.00 0.54 -10.78* -12.2** 3.47 4.63 5.07 25.00** 9.35* 11.2* 3.47 6.23 5.85 25.54** -6.15 28.2** 3.47 4.37 4.54 15.73** 3.89 -0.05 4.30 4.20 4.54 6.87 5.58 -0.05 4.30 4.48 4.07 -7.25* -9.14* -10.74* 4.30 4.63 4.92 10.07** 6.12 7.89 4.30 6.23 5.81 10.38** 35.19** 27.41** 4.30 4.37 4.61 6.46 6.65 1.10 4.20 4.48 4.01 -7.53* -10.45* 12.06** 4.20 4.63 4.45 0.72 -4.03 -2.41 4.20 6.23 6.25 19.85** 0.27 37.1** 4.20 4.37 4.44 3.62** 1.60 -2.63 4.48 4.63 4.97 9.10* 7.34 8.99* 4.48 6.23 5.62 4.82 25.28** 23.2** 4.48 4.37 4.58 3.58 4.96 0.44 4.63 6.23 6.23 14.66** 34.46** 36.6** 4.63 4.37 4.78 6.3 3.24 4.82 6.23 4.37 6.37 20.13** 2.14 39.7** 4.56 0.19 0.17 0.21 0.21 0.37 0.33 0.41 0.41 0.48 0.44 0.54 0.55 Fruit yield (g/plant) Per se performance Per cent heterosis Female Male F1 MP BP CC 1010.0 763.33 1333.33 50.38** 32.01** 37.02** 1010.0 623.33 866.67 6.12 -14.19** -10.92* 1010.0 983.0 1288.6 29.32** -27.59** 32.44** 1010.0 1131.67 866.67 -19.07** -14.19** -10.92* 1010.0 736.67 953.33 9.16 -5.61 -2.02 1010.0 900.00 996.67 4.36 -1.32 2.43 763.33 623.33 1058.67 52.69** 38.69** 8.80 763.33 983.0 926.67 6.13 -5.73 -4.76 763.33 1131.67 708.33 -25.24** -37.00** -2.72 763.33 736.67 823.33 9.78 7.86 -15.38* 763.33 900.00 1003.33 20.64** 11.48* 3.11 623.33 983.0 1045.00 30.11** 6.31 7.39 623.33 1131.67 1176.67 34.09** 3.98 20.92* 623.33 736.67 653.33 -3.92 -11.31 -32.85** 623.33 900.00 643.3 -15.54** -28.52** -33.88** 983.0 1131.67 880.00 -16.77** -22.24** -9.55 983.0 736.67 691.67 -19.56** -29.64** -28.9** 983.0 900.00 1005.0 6.54 2.24 5.21 1131.67 736.67 680.00 -27.21** -39.91** -30.11** 1131.67 900.00 800.00 -21.25** -29.31** -17.78** 736.67 900.00 933.33 14.05* 3.70 -4.08 973.30 48.09 42.35 48.90 48.90 94.25 83.00 95.84 95.84 123.59 108.8 125.67 125.69

Crosses S-22 x L-15 S-22 x CO-3 S-22 x Sivap S-22 x Solan Vajra S-22 x Local S-22 x PKM-1 L-15 x CO-3 L-15 x Sivap L-15 x Solan Vajra L-15 x Local L-15 x PKM-1 CO-3 x Sivap CO-3 x Solan Vajra CO-3 x Local CO-3 x PKM-1 Sivap x Solan Vajra Sivap x Local Sivap x PKM-1 Solan Vajra x Local Solan Vajra x PKM-1 Local x PKM-1 MHTM-256(CC) S.Em± CD at 5% CD at 1%

*Significant at 5% level of probability **Significant at 1% level of probability MP ­ Mid parent, BP-Better parent, CC-Commercial check

4.2.2.18 Fruit yield (g) The data on per se performance of parents and F1s (Table 30) revealed that the parent Solan Vajra (1131.67 g) had given highest fruit yield per plant and CO-3 (623.33) the least and the hybrid S-22 x L-15 recorded highest yield of 1333.3 while lowest by CO-3 x PKM-1 (643.3 g). The data on per cent heterosis revealed that the hybrids exhibited per cent heterosis in the range of -27.21 (Solan Vajra x Local) to 52.68 (L-15 x CO-3), -37.00 (L-15 x Solan Vajra) to 38.69 (L-15 x CO-3) and -33.88 (CO-3 x PKM-1) to 37.02 (S-22 x L-15) per cent respectively over mid parent, better parent and commercial check. Out of 21 hybrids, seven, three and three hybrids exhibited significant positive average heterosis, heterbeltiosis and standard heterosis, respectively. Among 21 hybrids, the cross S-22 x L-15 had highest positive heterosis of 50.38 per cent over mid parent, 32.01 per cent over better parent and 37.02 per cent increased vigour over the commercial check hybrid (MHTM-256).

4.2.3 Combing ability

4.2.3.1 Combining ability variance Analysis of variance for combining ability with respect to 18 characters are presented in the Table 31. From this analysis it was evident that both GCA and SCA variances were significant for all character thus indicating the importance of both additive and non-additive gene action in the inheritance of these characters. 4.2.3.2 Combining ability effects The general combining ability effects (gca) and specific combining ability effects (sca) were estimated for seven parents and 7 x 7 half diallel crosses without reciprocals respectively. The estimates for all the 18 characters including yield and yield component traits are presented in Tables 32 and 33 and the results are given below. 4.2.3.2.1 Plant height Among the seven parents studied, the parents L-15 (7.00), Local type (3.55), Solan Vajra (2.84) and Sivap (1.49) had significant positive gca effects while, PKM-1 (-2.80), S-22 (3.61) and CO-3 (-8.47) showed significant negative gca effects for plant height. In general 4 genotypes showed significant positive gca effects and 3 genotypes showed significant negative gca effects. The highest positive sca effects were recorded by the crosses L-15 x PKM-1 (25.64), CO-3 x Solan Vajra (18.06) and Local x PKM-1 (14.08). Totally five crosses had significant positive sca effects and five crosses had significant negative sca effects. 4.2.3.2.2 Number of primary branches The genotype which recorded significant positive gca effects was Local type (0.87), while Sivap and PKM-1 (-0.18 each), S-22 (-0.21) and CO-3 (-0.39) had significant negative gca effects. Of 21 crosses, S-22 x Local (0.87), Local x PKM-1 (1.34) and L-15 x PKM-1 (1.33) showed positive sca effect, while the cross, Solan Vajra x PKM-1 showed significant negative sca effect. Out of 21 hybrids, 9 recorded significant sca effects of which three recorded positive and 6 recorded significant negative sca effects. 4.2.3.2.3 Number of flowers per truss Among the seven parents, Local type (1.00) and PKM-1 (0.19), exhibited significant positive gca effects while remaining viz., Solan Vajra (-0.41), Sivap (-0.39), L-15 (-0.26) and S-22 (-0.13) showed significant negative gca effects. The cross S-22 x L-15 exhibited highest positive significant sca effects (0.71). Out of 21 hybrids, 9 exhibited significant positive sca effects and 5 showed significant negative sca effects. The cross CO-3 x PKM-1 showed significant negative sca effects (-1.16).

Table 31. Analysis of variance for combining ability in tomato Sl. No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Source Plant height (cm) Number of primary branches Number of flowers per truss Number of fruits per truss Number of fruits per plant Average fruit weight (g) Number of locules per fruit Fruit shape index (L/D) Pericarp thickness (mm) Ascorbic acid (mg/100 g) Total acidity (%) Reducing sugar (%) Non-reducing sugar (%) Total sugars (%) Juice recovery (%) Pulp content (%) Total soluble solids (°Brix) Yield per plant (g) Mean sum of squares SCA 152.06** 0.569* 0.341* 0.187* 25.82** 24.87** 0.231* 0.0033* 0.140* 15.756** 0.00264* 0.0545* 0.0141* 0.022* 31.939** 28.340** 0.140* 305.49** ²g 0.0064 1.049 0.0023 0.0026 0.263 0.0208 0.0136 0.0000489 0.017 0.0512 0.000008 0.000164 0.000105 0.000056 0.2279 0.226 0.0017 110.13 ²s 0.0546 8.873 0.0195 0.0236 2.222 0.1765 0.1153 0.000413 0.144 0.433 0.000072 0.00139 0.000894 0.000477 1.928 1.912 0.0144 931.55 ²g/²s 0.1172 0.1216 0.1179 0.101 0.1183 0.1178 0.1180 0.11840 0.11805 0.1182 0.1111 0.11798 0.11740 0.1174 0.1182 0.1895 0.11805 0.1200

GCA 246.85** 1.544* 2.169* 2.261* 348.31** 367.79** 1.696* 0.0203* 2.274* 43.464** 0.0011* 0.0570* 0.0251* 0.0099* 51.760** 47.415** 2.274* 535.38**

Error 11.015 0.0677 0.024 0.029 2.76 0.219 0.143 0.00051 0.0178 0.538 0.0009 0.00172 0.0011 0.00059 2.393 2.373 0.0178 115.64

*Significant at 5% level of probability **Significant at 1% level of probability GCA ­ General combining ability SCA ­ Specific combining ability ²g ­ Variance due to GCA = GA ²s ­ Variance due to GCA = VD

Table 32. General combining ability effects of seven parents for eighteen quantitative and quality traits in tomato

Sl. No. Parents Plant height Number of primary branches -0.21** Number of flower per truss -0.13** Number of fruits per truss -0.24** Number of fruits per plant -3.62** Average of fruit weight Number of locules per fruit 0.72** Fruits shape index Pericarp thickness Ascorbic acid Total acidity Reducing sugar Nonreducing sugar Total sugar Juice recovery Pulp content Total soluble solids Yield per plant

1

S-22

3.61** 7.00** 8.47** 1.49*

3.37**

0.04** 0.04** 0.04**

-0.36**

-3.18**

-0.02**

0.16**

-0.11**

0.05**

4.16**

-3.49**

-0.37**

116.08**

2 3

L-15 CO-3

0.05 -0.39**

-0.26** 0.00

-0.17** -0.19**

-6.47** 10.21** -5.73**

1.38* 0.11

-0.29** 0.09

-0.20** -0.25**

-0.76** 1.57**

0.02** 0.00

0.02* -0.03**

-0.01 0.06**

0.02** 0.03**

-2.01** -0.34

2.64** -0.07

-0.2** -0.24**

11.08* -65.58**

4

Sivap

-0.18**

-0.39**

0.3**

2.7**

0.07

0.04** 0.00

-0.39**

-0.24

0.00

-0.09**

0.04**

-0.05**

0.16

-0.9**

-0.29**

58.05**

5

Solan Vajra Local

2.84**

0.04

-0.41**

-0.22**

11.88** 44.16**

4.24**

-0.08

0.12**

0.24

-0.01**

-0.06**

0.01

-0.03**

0.35

-0.41

0.10**

10.49*

6

3.55**

0.87**

1.00**

1.11**

-14.18**

-0.69**

0.07**

-1.09**

3.73**

0.00

0.03**

-0.01

0.00

-3.42**

3.24**

1.09**

119.14** -10.38*

7

PKM-1

2.80** 0.68 1.33

-0.18**

0.19**

0.01

-6.25**

2.38*

0.17*

0.06** 0.005 0.01

-0.08**

-1.36**

0.00

-0.02*

0.01

-0.01*

1.09**

-1.00**

-0.08**

S.Em± CD at 5% CD at 1%

0.053 0.11

0.032 0.063

0.035 0.069

0.34 0.66

0.955 1.87

0.077 0.15

0.03 0.059

0.15 0.29

0.002 0.004

0.0083 0.017

0.0067 0.013

0.0049 0.01

0.314 0.61

0.313 0.60

0.03 0.09

6.91 8.82

1.75

0.14

0.082

0.090

0.87

2.46

0.20

0.013

0.077

0.38

0.0054

0.023

0.017

0.014

0.80

0.80

0.077

11.60

*Significant at 5% level of probability **Significant at 1% level of probability

Table 33. Specific combining ability effects for eighteen quantitative and quality characters in F1 hybrids of tomato

Sl. No. Parents Plant height Number of primary branches -0.64* -0.3 -0.65* -0.13 0.87** 0.27 0.17 0.09 -0.36 -0.31 1.33** 0.35 0.44 -0.96** -0.65* -0.24 0.51 -0.35 -0.65* -1.38** 1.34** 0.265 0.51 0.768 Number of flower per truss 0.71** 0.12 -0.22 0.29 0.34* -0.28 0.6** -0.34* 0.62** -0.67** -0.12 0.46* -0.54** 0.34* -1.16** -0.06 0.53** 0.01 0.51** -0.61** 0.61** 0.16 0.33 0.41 Number of fruits per truss 0.35* -0.19 -0.22 -0.12 0.23 -0.09 0.4* -0.15 0.25 -0.05 -0.23 0.12 -0.06 0.37* -0.49** -0.24 0.68** -0.03 0.61** -0.06 0.63** 0.17 0.33 0.44 Number of fruits per plant 2.02 -6.57** 0.61 -3.57* 31.39** -8.87** 6.61** -1.54 -3.72* 14.24** 10.02** 5.87** 7.35** -8.35** -4.28* -6.8** 15.17** -4.09* -5.02** -2.94 38.02** 1.7 3.3 4.38 Average of fruit weight 3.77** 2.54** 1.76** -2.96** -5.24** 2.74** 1.44** -2.01** -3.66** -2.54** 9.93** 0.02 5.81** -1.02* -4.07** 4.46** -5.71** 1.15* -5.08** -2.33** -4.68** 0.47 0.92 1.21 Total soluble solids 0.22 0.32* -0.19 0.49** 0.29* 0.14 0.13 -0.3* 0.16 0.07 0.04 -0.31 -0.26 0.55** -0.09 0.31* -0.03 0.1 0.2 -0.08 0.51** 0.137 0.27 0.35 Number of locules per fruit -0.09 0.47 0.31 -0.43 -0.8* 0.42 -0.23 0.90* 0.53 0.11 0.12 -0.16 0.11 -0.40 0.10 0.36 -0.39 -1.04** -0.07 0.18 -0.08 0.39 0.76 1.00 Fruits shape index 0.03 0.00 0.00 -0.02 -0.02 0.00 0.01 -0.01 -0.04 0.00 0.08** 0.00 0.02 0.05* 0.08** 0.08** 0.16** -0.01 -0.03 0.06* 0.01 0.023 0.045 0.059 Pericarp thickness Ascorbic acid Total acidity Reducing sugar Nonreducing sugar -0.05 0.11** 0.05 -0.09** -0.01 -0.02 -0.05 0.02 0.12** 0.14** 0.08* 0.14** 0.03 -0.17 -0.26 0.16 0.0 -0.05 0.02 -0.02 0.05 0.033 0.064 0.085 Total sugar Juice recovery Pulp content Yield per plant 296.09** -93.91** 204.46** 169.98** 46.31 -18.5 203.0** -52.54 -223.3** 21.3 93.17** 142.4** 321.7** -72.1* -190.1** -98.6** -157.3** 47.87 121.43** 109.57** 153.39** 34.84 68.28 89.88

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21

S-22xL-15 S-22xCO-3 S-22xSivap S-22xSolan Vajra S-22xLocal S-22xPKM-1 L-15xCO-3 L-15xSivap L-15xSolan Vajra L-15xLocal L-15xPKM-1 CO-3xSivap CO-3xSolan Vajra CO-3xLocal CO-3xPKM-1 SivapxSolan Vajra SivapxLocal SivapxPKM1 Solan VajraxLocal Solan VajraxPKM-1 LocalxPKM-1 S.Em± CD at 5% CD at 1%

12.71** 8.26* 2.7 6.52 5.33 -2.58 3.07 -3.32 2.34 3.30 25.64** 9.44* 18.06** 8.52** -25.6** -14.7** 6.56 2.82 -6.96* -7.2* 14.08** 3.4 6.66 0.77

0.22 0.10 0.16 0.30* -0.13 0.14 -0.10 -0.40** 0.10 -0.50** -0.56** -0.39 -0.29 0.11 -0.11 0.26 -0.62** -0.41** -0.36** -0.23 -0.36* 0.133 0.277 0.343

-2.44** -4.6** 1.79* -6.01** 0.74 3.24** -1.67* 3.12** -1.62** 4.49** -0.09 6.05** 6.16** -2.59** -1.34 2.55** -0.19 -0.19 -0.05 2.05* 5.86** 0.75 1.47 1.93

0.04** 0.00 0.01 0.07** 0.00 0.10** 0.01 0.00 0.06** 0.04** 0.00 0.06** 0.09** -0.02* 0.10** -0.02* 0.03* -0.02* 0.03* -0.02* 0.02* 0.009 0.017 0.023

0.28** -0.32** -0.21** -0.06 0.15** 0.1* 0.01 -0.16** -0.111* -0.39** -0.14** -0.13** 0.08 0.32** 0.43** -0.08 0.00 0.11** -0.18** -0.19** 0.08 0.042 0.088 0.108

0.24** 0.22** 0.15** 0.17** 0.14** 0.11** -0.05 0.14** 0.02 -0.23 -0.05 0.03 0.12** 0.17** 0.15** 0.09** 0.02 0.07** 0.13** 0.21** 0.12** 0.025 0.049 0.064

8.14** 5.42** -0.98 3.9* 5.64** -2.71 -2.01 -8.17** 4.91** 8.68** -1.07 3.46* -3.13* -2.26 4.16* 1.70 -4.26* 2.93 -7.63** -5.39** -9.72** 1.58 3.09 4.07

5.62** -0.64 1.10 -0.83 -8.85** 7.34** 1.09 7.47** -6.04** 2.20 0.14 -1.56 3.66* 1.29 -3.93* -2.42 5.29** -2.03 6.57** 4.18** 7.41** 1.57 3.07 4.05

4.2.3.2.4 Number of fruits per truss Of the seven parents only one recorded significant positive gca effect while five showed significant negative gca effect. The highest positive effect was recorded by a lone parent Local type (1.11) while, the highest negative effect by Sivap (-0.3). Among 21 hybrids, six exhibited significant positive gca effect and two showed significant negative gca effects. 4.2.3.2.5 Number of fruits per plant The genotypes that showed significant positive gca effect was Local type (44.16) whereas rest of the parents showed significant negative gca effect. The genotype Solan Vajra showed highest negative effect of -11.88. Of 21 combinations, the crosses Local x PKM-1 (38.02), S-22 x Local (31.39), Sivap x Local (15.17) recorded maximum positive sca effect while the crosses L-15 x PKM-1 (-10.02), S-22 x PKM-1 (-8.87) and CO-3 x Local (-8.33) showed maximum negative sca effect. Totally seven genotypes showed significant positive effect and 12 showed significant negative effect. 4.2.3.2.6 Average fruit weight Out of seven parental genotypes six showed significant positive gca effect and the lone parent showing significant negative gca effect was Local type (-14.18). The parent Solan Vajra exhibited highest significant positive gca effect (4.24). Among the hybrids, nine showed significant positive sca effect, of which L-15 x PKM1 recorded maximum positive sca effect (9.93). Eleven genotypes exhibited significant negative sca effect of which the cross, Sivap x Local (5.71) exhibited highest negative sca effect. 4.2.3.2.7 Total soluble solids Except two parental genotypes, Solan Vajra (0.10) and Local type (1.09) showed significant positive gca effects, while the rest showed significant negative sca effect of which S-22 (-0.37) exhibited highest negative sca effect. Though 15 crosses exhibited positive sca effect, only six crosses showed significant positive sca effect. The cross with negative and significant sca effect was L-15 x Sivap (0.13). 4.2.3.2.8 Number of locules per fruit Out of seven parents, four showed significant gca effect of which S-22 (0.72) and PKM-1 (0.17) showed positive gca effects and L-15 (-0.29) and Local type (-0.69) negative gca effects. Out of 21 hybrids evaluated only 3 exhibited significant sca effects, of which the cross L-15 x Sivap (0.9) exhibited positive sca effect and Sivap x PKM-1 (-1.04) and S-22 x Local (0.8) showed negative sca effect. 4.2.3.2.9 Fruit shape index The gca effects of parents L-15 (0.04), CO-3 (0.04) and Local type (0.07) were found to be significantly high and positive in direction. Of the remaining four parents S-22, Sivap and PKM-1 had negative significant gca effects, while, PKM-1 (-0.06) showed highest gca effects. Though 50 per cent of the hybrids (12) showed positive sca effects, only three registered significant positive sca effects and the cross, Sivap x Local (0.16) had highest positive sca effect. In the rest of the hybrids (11), only three registered significant negative sca effect and those were L-15 x PKM-1 (-0.08), CO-3 x PKM-1 (-0.08) and Sivap x Solan Vajra (-0.08). 4.2.3.2.10 Pericarp thickness All the parents, except Solan Vajra (0.12), showed significant negative gca effect, of which the parent Local (-1.09) was very poor general combiner for the pericarp thickness.

A total of eight hybrids showed significant sca effect for pericarp thickness, out of which only one cross S-22 x Solan Vajra (0.3) showed significant positive specific combining ability effect and in the rest of hybrids which showed significant negative sca effect the cross Sivap x Local (-0.62) registered highest negative gca effect. 4.2.3.2.11 Ascorbic acid The positive significant gca effects were observed in only three parents while negative gca effects observed in four parents. Out of seven parents, Local type (3.73) and CO-3 (1.57) were good general combiners which exhibited positive gca effect while the parent S-22 (-3.18) was poor general combiner for ascorbic acid. A total of 15 hybrids showed significant sca effect for ascorbic acid. Out of which eight crosses exhibited significant positive sca effect and highest was observed in CO-3 x Solan Vajra (6.16), CO-3 x Sivap (6.05) and Local x PKM-1 (5.86). Seven hybrids showed significant negative sca effect. The cross S-22 x Solan Vajra (-6.01) was a poor combiner for flesh thickness. 4.2.3.2.12 Total acidity Except three parents (S-22, L-15 and Solan Vajra) all the remaining exhibited nonsignificant gca effect. Only L-15 showed significant positive gca effect and the parents S-22 (0.02) and Solan Vajra (-0.01) showed significant negative gca effect. The significant specific combining ability effects were observed for 15 hybrids of which 7 exhibited positive and the remaining exhibited negative effects for the total acidity. Out of seven crosses S-22 x PKM-1 (0.10) had maximum sca effects followed by CO-3 x Sivap and L-15 x Solan Vajra which are on par with each other for sca effect (0.06). Two crosses (CO-3 x Solan Vajra and S-22 x Solan Vajra) exhibited significant negative sca effects. 4.2.3.2.13 Reducing sugar Among the 7 parents, three exhibited significant positive gca effect of which S-22 (0.16) recorded highest gca effect. Four parents showed significant negative gca effect of which the parent, Sivap (-0.09) registered highest negative gca effect. Though, 15 hybrids out of 21 showed significant sca effect for reducing sugar, only five hybrids were positively significant. CO-3 x PKM-1 (0.43) hybrid recorded maximum specific combining ability effect. The cross showing highest negative and significant sca effect was L-15 x Local (-0.39). 4.2.3.2.14 Non-reducing sugar Except two parents CO-3 and Sivap that showed significant positive gca effects, all other parents recorded non-significant gca effects. CO-3 and Sivap parents registered gca effect of 0.06 and 0.04 respectively. Though, 11 crosses exhibited positive sca effect only five had significant and positive sca effect. Out of them the crosses CO-3 x Sivap and L-15 x Local (both of them with sca effect of 0.14). L-15 x Solan Vajra and S-22 x CO-3 (0.11) showed maximum positive sca effect in the order of their merit. Out of nine having negative sca effect only one, S-22 x Solan Vajra (-0.09) showed significant sca effect. 4.2.3.2.15 Total sugar Out of 7 parents except one rest of them showed significant gca effect. Fifty per cent of the parents showed significant positive gca effect. The highest positive gca effect was observed in the parent S-22 (0.05). While, the parent, Sivap (-0.05) showed highest negative gca effect. Out of 21 hybrids under study 15 showed significant sca effect of which only nine showed positive sca affect. The highest positive sca effect in the order of merit was recorded by the crosses viz., S-22 x L-15 (0.24), CO-3 x Local (0.17) and CO-3 x PKM-1 (0.15). Similarly the highest negative sca effects were exhibited by the crosses, S-22 x CO-3 (-0.22), Solan Vajra x PKM-1 (0.21) and S-22 x Solan Vajra (-0.17).

4.2.3.2.16 Juice recovery percentage The gca effects of the parents S-22 (4.16) and PKM-1 (1.09) were found to be significantly high and positive in the direction, whereas the parents Local (-3.42) and L-15 (2.01) showed significant and negative gca effect. Significant specific combining ability effects were observed for 14 crosses. Out of these, 8 crosses exhibited positive while 6 crosses exhibited negative significant effects respectively. The 3 best combinations for the juice recovery percentage in the order of merit were L-15 x Local (8.68), S-22 x L-15 (8.14) and S-22 x Local (5.64) with positive sca effects. The cross Local x PKM (-9.72) exhibited significant negative sca effect for the trait. 4.2.3.2.17 Pulp content (%) The positive significant gca effects were observed in only 2 parents and negative gca effects observed in three parents. Out of seven parents, Local type (3.24) and L-15 (2.64) were good general combiners exhibiting positive gca effects and the parent S-22 (-3.49) was poor general combiner for pulp content. Out of 11 crosses showing significant sca effects, eight crosses showed positive and three crosses recorded negative significant sca effects. The 3 best combinations for the pulp content in the order of merit were L-15 x Sivap (7.47), Local x PKM-1 (7.41) and S-22 x PKM-1 (7.34) with positive sca effects. The cross S-22 x Local (-8.83) exhibited significant negative sca effects for the trait. 4.2.3.2.18 Yield per plant Of the seven parents 4 exhibited significant positive gca effects and the rest significant negative gca effects. The gca effect was 116.08, 58.05, 11.38 and 10.49 for S-22, Sivap, L-15 and Solan Vajra respectively. The highest negative gca effect was exhibited by the parent, Local type (-119.14) followed by CO-3 (58.05). Out of 21 crosses evaluated seven recorded significant positive sca effect and nine crosses recorded significant negative sca effect. The hybrid CO-3 x Solan Vajra (321.7) recorded the highest positive significant sca effects followed by S-22 x L-15 (296.09), S-22 x Sivap (204.46), L-15 x CO-3 (203.09 and Local x PKM-1 (153.39). The cross L-15 x Solan Vajra (-223.3) recorded highest significant negative sca effect.

V. DISCUSSION

Tomato (Lycopersicon esculentum Mill.) is one of the most common and commercially important vegetable crops grown intensively in India. The yield and quality enhancement is a continuous process carried out by plant breeders. It requires the evaluation of genotypes and also creation of variability and breakdown of unfavourable associations. Therefore an attempt was made to enhance variability through biparental mating and to assess magnitude of heterosis and to estimate combining ability. The results obtained are discussed under the headings given below. 5.1 Biparental mating 5.2 Diallel mating

5.1 BIPARENTAL MATING

Genetic diversity or variability is the most essential requirement for any successful crop improvement programme. Most of the genetic variability available today in plant collections is the result of spontaneous mutation, recombination and exposure to natural selection. Cultivated tomato (Lycopersicon esculentum Mill.), being a self pollinated crop has inherent problem of epistatic interactions and linkage of desirable and undesirable characters which inherit as blocks of genes through generations. This makes it difficult to realize recombination of desired characters from simple pedigree or bulk method of breeding. Hence there is a need to break these undesirable linkages through deliberate shuffling of genes through recombination to increase the efficiencies of breeding schemes. The best way by which this can be realized is by following biparental mating in early segregating generations. There are several success stories in literature, especially in crops like wheat, safflower etc. (Yunus and Paroda, 1983 and Parameshwarappa et al., 1997) where biparental or inter mating was used for producing sufficient variability on which selection could be employed to realize superior recombinants. This would also retain variability during subsequent selfing generations until the objective of isolating a rare desired recombinant is achieved. The varieties developed by inter mating are expected to be as competent as that of hybrid and infact would have an edge over hybrid as varietal seeds are cheaper. As already mentioned, no report on such attempts is available on tomato. Keeping this in view, it was planned to advance the F1s viz., MHTM-256 and S-4-14, both commercial hybrids to F2. The biparental mating, among selected plants within F2 which was a normal process and a new approach of crossing selected plants of F2 of a population with plants of other F2 population was carried out. The comparison was made between the intra and intermating population and also between BIPs and F3, to test the hypothesis that inter mating in the early segregating generations like F2 of the cross, would provide increased opportunity for extra recombination and consequently the release of concealed variability from which there would be better scope to identify and select the desirable recombinants.

5.1.1 Mean

The comparison of mean performance of different characters between intra and inter mating populations (Table 7) indicated that in general intra population mating is better for all the characters like plant height, primary branches, fruits per plant, average fruit weight, locules per fruit, fruit shape index and fruit yield except fruits per truss and pericarp thickness. The characters, fruit yield, primary branches per plant, locules per fruit and fruit shape index recorded higher mean in intra mating population than inter mating populations. The comparison of mean performance of different characters between BIPs and F3 population indicated that, for the characters like primary branches, fruit/truss, locules per fruits fruit shape index and BIPs showed higher mean values compared to F3 but for the characters like plant height, fruits per plant, average fruit weight, pericarp thickness and yield per plant the F3 showed higher mean values than BIPs. It was due to the wider range observed in BIPs for all the characters understudy. Higher mean values in BIP compared to selfing generation was reported by Randhawa and Gill (1978), Yunus and Paroda (1998) and Singh et al. (1988) for traits like plant height, tillers per plant, 100-grain weight and grain yield in wheat and Singh and Sahu (1981) for number of branches per plant, number of capitula, 100-seed weight, grain yield in safflower.

Plate 1. Fruit shape and size variation observed in populations developed through intra mating

Plate 2. Fruit shape and size variation observed in populations developed through selfing

Plate 3. Fruit shape and size variation observed in populations developed through intermating

Plate 4. Superior segregant for fruit yield observed in MHTM-256 X S-4-14 population (3.34 kg/plant)

Among BIP populations, population S recorded higher mean values for the traits, plant height, primary branches, average fruit weight, locules per fruit, pericarp thickness and fruit yield, whereas population M/S showed higher mean value for traits like fruits per truss and fruit per plant. All BIP populations showed lower mean values for plant height, fruits per truss, fruits per plant and yield per plant compared to both of the commercial checks MHTM-256 and Numdhari hybrid. This can be attributed to the wide range of variability shown by the BIP populations for these traits. For the traits like primary branches, average fruit weight, locules per fruit and pericarp thickness BIPs showed better mean values than the commercial checks.

5.1.2 Range

It was already discussed that the mean for all of the traits except fruits per truss was higher in intra mating population compared to inter mating population. This is because, the range for most of the traits viz., plant height, fruits per truss, fruits per plant, average fruit weight, locules per fruit, fruit shape index and yield per plant (Table 8) was higher in case of inter mating populations than intra populations. Range for pericarp thickness is on par in all the populations of BIP. It is noteworthy that though the lower limit in general for all of the characters does not differed among the population but upper limit for the characters, fruits per plant, average fruit weight and yield per plant was far higher in inter mating populations compared to intra mating population. This suggests that inter mating has helped in releasing more variability. The higher variability in inter mating population could also result from the additional opportunity for genetic recombination. The range observed for most of the traits in BIPs was wider. It is noteworthy that especially the upper limit of range was higher in BIPs for all characters. At the same time the lower limit was smaller compared to that of F3 and commercial checks, suggesting intermating has helped in releasing more variability than in selfing generations. General shift in value of ranges of characters by biparnetal approach was also reported by Nemtullah and Jha (1993) in wheat and Parameshwarappa et al. (1997) in safflower. Among BIP populations, the population M/S showed wider range for characters fruits per plant and fruit yield. Infact one plant in this population showed yield which is almost double the upper limit values of all other populations. For characters like plant height, fruits per truss, average fruit weight, locules per fruit and fruit shape index the population S/M showed wider range compared to rest of populations. From above discussion we can deduce that based on mean value one should not reject a progeny or family. For example the mean value of population M/S for fruit yield was lowest compared to other segregating populations but the range for this character is wider so it gives scope for selecting superior segregants. It is also supported by taking another example of population S/M, which showed lower mean value for fruits per plant and locules per fruit but quite a wider range for these characters.

5.1.3 Variability

The total variability was divided into genotypic and henotypic variances, GCV and PCV, heritability and genetic advance. Genotypic and phenotypic co-efficient of variation The comparison of GCV and PCV in intra mating and inter mating population for nine traits (Table 9) indicated that the estimate of PCV were generally higher than GCV for all of the characters. This may be due to the involvement of high environmental and genotypic x environmental interaction effect in character expression (Kaushik et al. 1996). The characters which showed wider range were also characterized by higher magnitudes of GCV and PCV. Among BIPs, though intra mating population showed high GCV and PCV for the traits like primary branches, fruits per truss, locules per fruit, fruit shape index and pericarp thickness but for the trait of interest for breeders, yield per plant, the inter mating populations showed higher GCV and PCV compared to intra mating population. The other characters where the inter mating population showed higher GCV and PCV were average fruit weight and plant height. The GCV and PCV were comparable in both inter and intra mating population for the character fruits per plant.

Among the segregating populations in general, BIP population had higher GCV and PCV than selfs for all characters except fruits per plant yield per plant and fruit shape index. Higher GCV and PCV in BIPs as compared to selfs were also reported in wheat by Srivastava et al. (1989) and Nanda et al. (1990a) and Kadlera (1997) in safflower. Among the characters in BIPs, fruits per plant and primary branches showed high GCV and PCV followed by average fruit weight locules per fruit, fruits per truss, yield per plant, shape index, pericarp thickness and plant height. The high GCV and PCV in case of fruits per plant and primary branches can be attributed to the predominance of repulsion phase linkage for these traits. Randhawa and Gill (1978), Gurudev Singh et al. (1986), Shrivastava et al. (1984) and Kadlera (1997) reported high GCV and PCV for tillers and grain yield per plant and seed number, respectively. Kampli et al. (2002) also reported high PCV and GCV for plant height primary branches and seed weight in chickpea. This suggests that, there is more scope for selecting better segregants in BIP population on basis of fruits per plant and primary branches per plant. Comparison of different BIP populations for GCV and PCV reveals that, the population M/S showed highest GCV and PCV for the traits like yield per plant, average fruit weight and plant height and for the traits like fruits per truss, primary branches and fruits per plant the population M/S is second best as far as PCV and GCV is concerned. So selection in population M/S for important yield attributing traits like primary branches, fruits per plant fruits per truss and average fruit weight would result in better response. Heritability and genetic advance A comparison of heritability estimate between inter and intra mating populations revealed that, in general the intra mating population showed higher heritability in broad sense for characters like primary branches, fruits per truss, locules per fruit, fruit shape index and pericarp thickness. A comparable heritability in broad sense between inter and intra mating population for characters fruits per plant, average fruit weight and yield per plant was observed. But critical examination of all four populations revealed that the heritability in broad sense was in general better for the population M/S for the characters plant height, fruits per truss, fruits per plant, average fruit weight and yield per plant when compared to rest of the populations. Among the traits studied average fruit weight showed highest heritability estimate followed by yield per plant, fruits per plant, plant height, primary branches, locules per fruit, fruit shape index, pericarp thickness and fruits per truss. A comparison of heritability estimates between BIPs and the selfed population revealed that, heritability estimates improved considerably for almost all the characters in BIPs except for fruit shape index where heritability of population derived from inter mating showed less heritability than selfed population. The change of heritability estimates towards higher side in biparental progenies over selfing series occurred probably be due to increased proportion of genetic variance to total phenotypic variance due to cryptic genetic changes that have been brought about by one cycle of inter mating. This suggests that the variation in the environment played a relatively limited role in influencing the inheritance of these characters. Improvement in heritability values for yield and its component traits is of particular interest to the breeder as it enhances the scope for improved selection response for characters such as fruits per plant, average fruit weight and number of primary branches. High heritability estimates in case of BIPs as compared to selfed progenies were also reported by Yunus and Paroda (1983), Nanda et al. (1990) in wheat and Kadlera (1997) in safflower. Genetic advance also showed similar pattern as that of heritability in broad sense. BIP populations showed relatively high genetic advance as per cent mean estimates for all the characters compared to selfed generations. Among the characters average fruit weight showed highest genetic advance followed by primary branches. This suggested that gain from selection based on these two traits would be higher in BIP population than in their corresponding selfed progenies. Increase in genetic advance through inter mating was also reported by Srivastava et al. (1989), Nanda et al. (1990b) and Kadlera (1997) in wheat and safflower respectively. Sharma and Kalia (2003) in garden pea also reported similar results as above workers for pod yield per plant and pods per plant.

The genetic advance as percent mean was higher for inter mating population for the character of much interest for breeders that is yield per plant and also for plant height and average fruit weight. Though for other characters like primary branches, fruits per truss, fruits per plant, locules per fruit, fruit shape index and pericarp thickness the intra mating population showed higher GA as per cent of mean compared to inter mating populations, the population M/S, an inter mating population, showed higher GA as per cent of mean for characters plant height, fruits per truss, fruits per plant, average fruit weight and yield per plant.

5.1.4 Inter character correlations

The release of hidden variability can also be examined by studying the changes in correlation pattern in different segregating population. The nature of association among different traits (Table 10 to 14) have been discussed here keeping in view the shift in correlation coefficient in BIP populations and F3. In general the magnitude of correlation co-efficient in intra mating population increased compared to inter mating population, for example, correlation between fruit yield and primary branches, fruits per truss and number of fruits per plant, primary branches and number of fruits per plant. In general, magnitude of correlation co-efficient in segregating populations of BIP have increased compared to selfed population, for example, the correlation between yield per plant and plant height, primary branches and yield per plant, plant height and number of fruits per plant. The increase in magnitude of correlation co-efficient would be expected if linkages were in repulsion phase (Nanda et al., 1990b). However in all the populations, association of number of fruits per plant with fruit yield and primary branches with fruit yield was high, positive and significant. This indicates that fruits per plant and primary branches per plant are the most important yield contributing characters in tomato and therefore emphasis should be given for these traits while making selection. The association between average fruit weight with fruits per truss which was non significant in intra mating population changed to significant and negative in inter mating population. Among BIP populations the change in association was observed. For example, in population S/M the association of yield per plant with number of fruits per truss and average fruit weight which was insignificant changed to significant and positive in rest of the populations under study. Shift in the correlations has also been reported earlier in wheat by Yunus and Paroda (1982), Nanda et al. (1990b) between character seed yield and plant height, seed yield and 100 grain weight. In safflower Kadlera (1997) and Parameshwarappa et al. (1997) observed shift in correlations between characters plant height and seed yield, number of branches and seed yield and number of capitula per plant and seed yield.

5.1.5 Path coefficient analysis

Path coefficient analysis further provided an insight into the inter relationships of various characters with fruit yield. A comparison of the direct and indirect effects of various characters on fruit yield (Table 15 to 19) in various BIP and F3 populations revealed that the change in the nature and degree of association amongst various characters was accompanied by the change in their direct and indirect effects. For example, fruits per plant exhibited high and positive direct effect on fruit yield and it was also positively and significantly associated with fruit yield in all the populations. This suggests that fruits per plant is a major factor contributing to fruit yield which also observed in tomato by earlier worker (Padda et al. 1971, Nandpuri et al., 1973, 1976 and 1977, Nagaraj, 1981, Bhutani and Kalloo, 1989, Prasad and Prasad, 1997, Rathod, 1997 and Patil, 1998). In all the populations of BIP the indirect effect of number of branches on the yield per plant via number of fruits per plant was observed and it is in agreement with Patil (1998). In all the populations it was observed that the direct effect of number of fruits per plant was highest followed by average fruit weight and number of primary branches. Therefore, by selecting for more number of primary branches and fruits per plant with higher average fruit weight in BIPs, the fruit yield can be improved. It can also be observed that the locules per fruit had direct positive effect on yield but its effect was counter balanced mostly by indirect negative effect via number of fruits. Similar result was observed by Bhutani and Kalloo (1989). The indirect effect of fruits per truss on yield per plant via the number of fruits per plant was also observed. A change in path for the characters was observed in inter mating population compared to intra mating population. For example, the direct effect of plant height on fruit yield was negative in

intra mating population while positive in inter mating population and indirect effect of pericarp thickness via locules per fruit was negative in intra population but positive in inter mating population. A change in path for the characters was observed in BIPs compared to selfed progeny. For example, the indirect effect of fruits per truss via primary branches on fruit yield was negative in selfed progeny but it changed to positive in BIP population.

5.1.6 Identification of superior segregants for yield

The segregating population should be assessed using means and variability along with their ability to release superior segregants, to know the real worth of a population. In the present study, extent of superior segregants for yield was identified in segregating populations, for which the check MHTM-256 was considered as base and are presented in Table 20. When compared to intra mating, inter mating has resulted in realizing quite a high number of superior segregants. Infact a plant in population M/S recorded double the yield per plant than the best superior segregant in any of the populations studied. Among the different BIP populations studied, the population S/M recorded high number of superior segregants. Between BIPs and F3 the BIP showed higher number of superior segregants. The higher frequency of superior segregants in BIPs particularly in inter mating population might be mainly due to more chance of recombination between favorable genes and also due to accumulation of more number of favourable genes. The per se performance of individual superior segregants for the group of important traits viz., primary branches fruits per plant, average fruit weight and yield per plant in different segregating population is given in Table 34. As can be seen from the table, plant 16 in population M/S was the best segregant with yield of 3.340 kg/plant showing the average fruit weight, fruits per plant and primary branches of 42.8 g, 78 and 9 respectively. The next best was from population S/M viz., plant 318 with 1.65 kg yield per plant and an average fruit weight of 63.4 g with 7 primary branches. Interestingly both plants belong to the same inter mating population. The best segregant for average fruit weight was plant 318 of population S/M followed by plant 11 of population S with per se values of 63.4g and 58.8g respectively. For number of fruits per plant and primary branches the best segregant was plant 16 of population M/S. The inter mating population in general recorded higher number of primary branches than intra mating population. The inter mating population showed little improvement in fruit weight along with high number of fruits per plant whereas the intra mating population showed little higher average fruit weight but the number of fruits per plant was low compared to inter mating population. Therefore it may be concluded that the objective of improving yield by improvement in both average fruit weight and fruits per plants is better achieved by inter mating than by intra mating. Compared to F3, the BIP recorded high primary branches, fruits per plant. The reduction in yield compared to BIPs can be attributed to reduction in number of fruits per plant, plant height, primary branches and average fruit weight.

Table 34. Per se performance of individual superior segregants for nine characters in different BIP and F3 populations in tomato

Population Sl. No Population M 1 2 3 4 5 Population S 1 2 3 4 5 6 7 Population M/S 1 2 3 4 5 6 7 8 9 10 Population S/M 1 2 3 4 5 6 7 8 9 10 F3 1 2 3 4 5 Check ­ MHTM-256 Namdhari (F1) Plant no. 6 36 19 30 20 19 22 20 26 29 11 46 16 4 24 17 12 14 32 35 43 18 318 299 91 296 92 127 98 320 104 129 13 28 19 18 35 Plant height (cm) 60.00 65.00 55.00 45.00 55.00 70.00 70.00 65.00 60.00 55.00 70.00 60.00 100.00 90.00 65.00 60.00 60.00 100.00 100.00 60.00 55.00 80.00 70.00 60.00 70.00 60.00 80.00 60.00 60.00 55.00 70.00 105.00 65 60 65 55 55 73.30 70.1 Number of primary branches 8 5 5 6 6 6 7 6 7 7 6 6 9 7 6 7 6 8 6 5 6 7 7 7 6 8 6 8 7 7 5 7 7 7 6 6 6 3.66 3.7 Fruits per truss 3.1 4.1 3.1 3.1 3.3 2.0 2.1 4.0 2.3 3.2 2.3 3.2 3.5 3.4 2.2 3.3 2.5 2.3 3.6 2.3 2.3 3.1 2.1 3.2 4.4 3.3 3.1 3.3 2.3 3.4 3.1 3.3 2.2 3.1 2.8 2.3 2.5 3.33 3.2 Fruits per plant 29 33 20 30 47 30 25 32 23 25 18 22 78 36 32 38 37 45 39 29 37 40 26 40 38 30 28 25 23 30 28 52 28 29 28 36 35 31 30 Average fruit weight (g) 52.0 45.1 50.0 38.3 23.8 51.0 58.4 39.7 53.4 46.4 58.8 47.7 42.8 43.3 45.6 36.0 36.4 30.0 34.6 41.3 32.4 30.0 63.4 34.7 39.4 49.5 50.7 54.4 56.9 43.3 46.4 24.0 58.5 52.0 54.0 32.5 30.0 32.0 33.0 Locules per fruit 5.00 2.30 7.00 5.20 3.00 4.20 6.20 5.50 3.10 5.20 5.20 4.30 5.00 5.10 3.30 4.00 4.00 3.00 6.10 3.10 5.20 6.00 4.10 3.10 3.10 3.10 4.30 5.10 5.50 2.10 5.10 4.20 4.1 5.0 4.9 3.5 3.8 3.10 3.20 Shape Index 0.75 0.60 1.00 0.80 1.30 1.30 1.24 1.26 1.15 0.59 1.20 1.40 1.15 0.85 0.60 0.40 0.67 0.90 0.73 0.75 0.70 0.40 1.26 1.36 1.25 1.22 1.17 1.24 0.92 0.89 0.93 0.80 1.2 0.8 0.9 0.9 0.6 0.91 0.80 Pericarp thickness (cm) 0.30 0.30 0.40 0.40 0.30 0.40 0.40 0.40 0.40 0.40 0.50 0.30 0.30 0.40 0.30 0.90 0.40 0.30 0.50 0.30 0.40 0.73 0.40 0.40 0.40 0.50 0.50 0.40 0.40 0.50 0.40 0.30 0.3 0.3 0.4 0.4 0.3 0.34 0.36 Yield per plant (g) 1510 1450 1200 1150 1120 1530 1460 1270 1230 1160 1060 1050 3340 1560 1460 1370 1350 1350 1350 1200 1200 1200 1650 1600 1500 1490 1420 1360 1310 1300 1300 1250 1650 1500 1500 1180 1140 973.3 988.4

5.2 DIALLEL MATING

One of the most perplexing challenges to agricultural research is the ways and means of achieving food and nutritional security. So far as nutritional security is concerned, production of sufficient quantity of food and good quality of vegetables and fruits holds the key. Year round availability, easy culture, high yield and consumption in varied ways (fresh or processed form) has made tomato as one of the most important vegetable crops of the world. However burgeoning population demands additional tomato production in large quantities. One of the inevitable ways to address this is by increasing the production. Any further delay in addressing this problem will result in accentuating to a point where nutritional security will get utterly out of hand. Therefore improvement of tomato productivity with high nutritional quality can play a significant role in the overall nutritional security. If the relevant high yielding varieties of the hybrids are developed and made available to farmers, the situation could considerably improve and per hectare yield could go up. Keeping these in view studies on heterosis and combining ability were undertaken on tomato to determine the extent to which heterosis is manifested in this crop for high yield and quality. The salient results of the investigation are discussed in this chapter.

5.2.1 Heterosis

The impulse of progress in crop improvement through plant breeding was propelled by a better understanding and an appropriate exploitation of heterosis. Heterosis is manifested through greater vigour of F1s over the parents resulting into higher yields through their component characters. The term signifies an increased or decreased vigour of F1s over mid parent or best parental values. The sector of success of F1 hybrids lies in the fact that all plants are exceptionally uniform in growth and development, better adaptability to changing and adverse ecological conditions and above all give high total yields. The ultimate choice of the parents to be used in a breeding programme is determined by per se performance and their behavior in hybrid combinations. Some idea on the usefulness of the parents may be obtained from their individual performance, particularly in respect of yield components. It is therefore, necessary to assess genetic potentiality of the parents in hybrid combinations through systematic combining abilities. Diallel method has been used in the present study for estimating combining abilities. The recent trend in tomato breeding has been towards development of hybrids to meet the specific uses (viz., earliness, growth habit, disease or pest resistance, fruit quality for processing fresh market etc.) coupled with high yield as it may be difficult to develop a hybrid having all the characters. However, it is reasonable to search one which can have maximum number of useful characters keeping yield as primary motto. 5.2.1.1 Growth parameters For plant height, it is interesting to note that all the 21 crosses exhibited positive and significant heterosis except (CO-3 x PKM-1) over the commercial check MHTM-256. Over the best parent majority of crosses showed significant heterosis and the same trend observed for mid parent heterosis. The parental range for plant height was 60.67 cm to 95.17 cm. The highest plant height was recorded by Solan Vajra. The plant height for hybrids ranged from 51.57 cm to 118.33 cm. Based on the F1 per se compared to their respective parental values, the hybrid L-15 x PKM-1 showed highest plant height followed by L-15 x Local and CO-3 x Solan Vajra. Per se values also suggest the prevalence of over dominance for the trait. Most of the 21 hybrids showed significant derivation from their mid parent values with respect to plant height. In the earlier studies also, heterosis was reported for plant height by Mishra and Khanna (1997), Fageria et al. (2001) and Thakur et al. (2004). Primary branches is another important growth parameter contributing for productivity. The parental lines ranged from 6.32 (Local type) to 4.33 (PKM- 1) in respect of number of primary branches. The standard check had 3.66 mean number of primary branches. Compared to this F1 hybrids exhibited mean number of primary branches ranging from 6.2 to 3.45 indicating that there was no much difference than the parental range. However, compared to the standard check Local x PKM-1, L-15 x CO-3 and S-22 x Local showed relatively high and significant positive standard heterosis. Heterosis was mainly due to dominance, in the latter two crosses and was due to over dominance in first cross. The

heterosis for this character was reported by Bhutani et al. (1973), Bhatt et al. (1999) and Tiwari and Lal (2004). 5.2.1.2 Yield and yield components Flowers per truss is another component trait that contributes for productivity. Higher number of flowers per truss is desired. The parental values ranged from 3.27 (Sivap) to 5.42 (Local types), while the range of the F1s was 3.39 (Sivap x Solan Vajra) to 6.05 (Local x PKM1). When the heterosis over mid parent and better parent was considered, as many as ten and four hybrids showed positive heterosis over mid parent and better parent respectively. When compared to standard check more than 50 per cent of hybrids showed positive heterosis and further the magnitude of their positive deviation was much higher compared to the magnitude of negative deviation. S-22 x Local, CO-3 x Local and Sivap x Local, Sivap x Local showed positive standard heterosis of more than 60 per cent. Heteoriss for flowers per truss was reported by Sundaram (1994), Tendulkar (1994) and Patil (1997). In the present study, the mean values of 15 hybrids in respect of fruits per truss was higher than the corresponding mean values of (2.73) the commercial check. The range of fruits per truss in F1 was 2.47 (S-22 X Sivap and Sivap x Solan Vajra) to 4.99 (Local X PKM1). The parental lines ranged from 2.55 (Sivap) to 4.23 (Local type), indicating that the fruits per truss was no much difference than the hybrid range. Nine and four out of 21 hybrids showed significant and positive heterosis over mid parent and better parent respectively and eight of the hybrids showed positive and significant heterosis over the commercial check. Heterosis for fruits per truss was observed by Pujar and Kale (1994), Patil (1997), Bhatt (1999) and Tiwari and Lal (2004). Fruits per plant and average fruit weight together form the most important closely related productivity parameters. The parental range for number of fruits per plant was 28.67 (CO-3) to 95 (Local type). The parents Local type, S-22 and Sivap had significantly higher fruit number than the standard check. Out of 21 hybrids about 50 per cent (13) hybrids had higher number of fruits than standard check hybrid. Wherever Local type was used as one of the parents such crosses showed higher number of fruits per plant since all have more or less similar size as that of Local which has small size fruits. Nine hybrids showed positive significant heterosis over mid parent and five and 13 crosses showed significant and positive heterosis over better parent and commercial check respectively. The magnitude of heterosis over commercial check for fruits per plant was high with the highest value of 308.77 per cent (Local x PKM-1). For this character negative and positive heterosis was observed by Yogananda (1977), Pujari and Kale (1994), Sundaram et al. (1994), Dharmatti (1995) and Thakur (2004). The hybrids in general had lower average fruit weight compared to commercial check. The mean average fruit weight ranged between 7.43 grams to 37.6 g. The Local type has small size fruits with low average weight wherever Local type used as one of the parent such crosses showed lower average fruit weight as their size is small. Hybrid L-15 x PKM-1 with average fruit weight of 37.6 g may be regarded as the best. The number of hybrids showing significant positive heterosis, over standard check were three while those showing the negative heterosis was higher. Heteorsis for this character reported by Fageria et al. (2001), Thakur (2004) and Tiwari and Lal (2004). Fruit yield per plant is the ultimate and most important trait. Overall mean of the hybrids was higher than the parental mean though it was lesser than that of check. The highest mean value for this trait shown by the hybrid S-22 x L-15 (1333.3 g) is higher than the best parent S-22 (1010 g) and so also over that of the check (973.3 grams). Only 3 hybrids showed significant positive heterosis over the commercial check. Two of them involved S-22 and CO 3 as one of the parents. The other two parents involved in those three hybrids being L-15 and Solan Vajra. The heterosis for this character observed by Bhatt et al. (1999), Fageria (2001) and Thakur (2004). 5.2.1.3 Quality parameters In tomato, quality parameters, namely, total soluble solids, ascorbic acid, total acidity, and sugars determine the quality related to taste, flavour and utility to processing or fresh market. A hybrid with low acid content is preferred for fresh market. However for processing

purpose, varieties with high acid content is desirable. Higher total soluble solids (TSS) and ascorbic acid content is desirable for higher recovery of processed products. The sugars make an important contribution to the flavors of tomato and among them reducing sugars contribute at least 50 per cent of the total soluble solids. The parental lines ranged from 3.47 (S-22) brix to 6.23 (Local) ° brix in respect of total soluble solids. The standard check had 4.56 °brix total soluble solids, compared to F1 hybrids which exhibited mean TSS ranging from 4.00 ° brix (S-22 x Sivap) to 6. 37 ° brix (Local x PKM­ 1) indicating that there was no much difference than the parental range however compared to the standard check, the hybrids Local x PKM­1, CO 3 x Local and Solan Vajra x Local showed relatively high significant heterosis. In these crosses heterosis was mainly due to over dominance. The heterosis for this character observed by Dharmatti (1995), Patil (1997), Makesh (2002) and Tiwari and Lal (2004). The hybrids in general had higher mean ascorbic acid content compared to parental mean and check. The mean ascorbic acid content ranged between 17.08 (S-22 x L-15) to 31.70 mg/100 g (Local x PKM-1). The hybrid L-15 X PKM­1 with mean ascorbic acid content (31.70 mg/100 g) is considered as the best and is the hybrid which showed highest significant positive heterosis for total soluble solids. All of hybrids showed significant positive heterosis over standard check. Similar results are reported by Patil (1988), Makesh et al. (2002) and Tiwari and Lal (2004). The parental range for total acidity was 0.27 per cent (S-22) to 0.43 (C03), which is higher than standard check MHTM-256 (0.26). However, it is denoted that L-15, CO-3 and Solan Vajra had significant high total acidity than standard check. The mean performance of the F1, hybrids was also in general better than the standard check with all the hybrids having numerically higher total acidity than standard check. All the hybrids except two under study showed positive significant heterosis over standard check for total acidity and magnitude of these heterosis values of high with the highest value of 69.20 per cent (S-22 x PKM-1). The heterosis for titratable acidity was observed by Patil (1988) and Makesh et al. (2002). Sugars are one of the contributors to the flavours. In the present study 3 types of sugars viz., reducing sugar, non reducing sugar and total sugar were studied. A critical examination of the data on these characters in the material studied reveals the following things. Though the mean of the hybrids in respect to all 3 types of sugars was lower than the corresponding mean values of the commercial check, still many of the hybrids exhibited significant higher mean values than commercial check. The crosses S-22 x L-15, S-22 x Local and CO-3 x PKM-1 exhibited significant positive and high heterosis over commercial check for reducing sugars, the crosses CO-3 X Sivap, Sivap X Solan Vajra, L-15 X Solan Vajra and L15 x Local exhibited higher positive heterosis over commercial check for non-reducing sugar and for total sugars S-22 x CO-3 and S-22 x L-15 exhibited high positive and significant heterosis. Seven and three hybrids over mid parent and better parent respectively for reducing sugars, nine and eight over mid parent and better parent respectively for nonreducing sugars and eight and five over mid parent and better parent in total sugars showed positive and significant heterosis. Fruits with lower number of locules are preferred for processing. In the present study the mean of the hybrids was on par with that of the check. None of the total 21 hybrids showed lower locules per fruits compared to the commercial check (3.11). The positive and negative heterosis for locules per fruit in tomato observed by Sundaram et al. (1994), Reddy and Reddy (1994), Dod et al. (1996) and Dharmatti (1995). With respect to fruit shape index the overall mean of the hybrids was just lower than that of check and also less than that of parental mean. The highest mean value was recorded by Sivap x Local (1.02) which is higher than best parent (CO-3) and check both showing fruit shape index of 0.91. Only two hybrids showed significant positive heterosis over the commercial check. The positive and negative heterosis was observed by Dundi (1991), Reddy and Reddy (1994) and Tendulkar (1994). With regard to pericarp thickness it should be as high as possible as it determines the keeping quality. In the present study the mean of the hybrids is more than that of the check, and again the range for this traits recorded by the hybrids indicated quite a high number of hybrids with high pericarp thickness. Seven hybrids studied showed significant and positive higher pericarp thickness above the commercial check, which ranged from 41.17 (S-22 x L-

15) to 17.64 (CO 3 x PKM ­1 and Solan Vajra x PKM­1). Heterosis for pericarp thickness also reported by Patil and Patil (1988), Makesh et al. (2002) and Tiwari and Lal (2004). In the fresh market the juicy tomato are preferred. The parental lines ranged from 27.92 (CO-3) to 39.5 (PKM-1) in respect of juice content. The standard check had 22.7 per cent mean juice content. Compared to this F1 hybrids exhibited mean juice content ranging from 20.67 (Solan Vajra x Local) to 40.67 (S-22 x CO-3). However compared to standard check S-22x CO-3, S-22 x Solan Vajra and S-22 x Local showed high significant and positive standard heterosis. In the first two hybrids heterosis was due to over-dominance and in third hybrid it was due to dominance. For processing purpose the hybrids should have high pulp content. The parental lines ranged from 58.1 (S-22) to 67.0 (L-15) in respect of pulp content. The F1 hybrids exhibited mean pulp content ranging from 57.86 per cent (S-22 X Local) to 76.61 per cent (Local x PKM-1). However compared to commercial check no hybrid exhibited heterosis in positive direction.

5.2.2 Combining ability effects and variances

The combining ability analysis gives an indication of the variance due to GCA and SCA, which represent a relative measure of additive and non-additive gene actions, respectively. It is an established fact that dominance is a component of non additive genetic variance. Breeders use these variance components to infer the gene action and to assess the genetic potentialities of the parents in hybrid combination. The ultimate choice of parents to be used in a breeding programme is determined by per se performance and their behavior in hybrid combination. Some idea on the usefullness of the parents may be obtained from their individual performance particularly in respect of yield components. It is therefore necessary to assess genetic potentiality of the parents in hybrid combination through systematic studies in relation to general and specific combining abilities. Diallel method has been used in the present study for estimating combining ability. 5.2.2.1 Analysis of variance Analysis of variance for combining ability (Table 31) reveals the presence of both additive and non additive gene action in all of the 18 characters studied, which was indicated by the significance of both the GCA and SCA variances. Therefore, improvement for these traits in the material studied can be achieved by few cycles of recurrent selection. The variance due to GCA was lower in magnitude than SCA for all characters. Further, the values of ²g to ²s ratio for all the traits supports the predominance of non additive gene effects in governing expression of all these characters. These results are encouraging from point of view of heterosis exploitation. 5.2.2.2 General combining ability (GCA effects) Among seven parents, the highest and significant gca effect for yield per plant (116.08) was observed in S-22 followed by Sivap (58.05). These parents also had maximum gca effects for average fruit weight, number of locules per fruit, reducing sugars, non-reducing sugar, total sugars and juice recovery percentage, where as L-15 exhibited significant gca effect in plant height, average fruit weight, ascorbic acid content, total acidity, sugars, pulp content. Though Local type exhibited negative gca effect for yield per plant, it showed positive and significant gca effects for plant height, number of primary branches, number of flowers and fruits per truss, number of fruits per plant TSS, shape index, pulp content and reducing sugars. As Local type showed superiority for most of the traits, gene pool developed by intermating in the segregating population inviting this parent is expected to offer maximum promise in genetically enhancing yield (Singh, 1982). For long term approach, the value of developing population may be emphasized and use of Jensons's (1970) method is suggested. The parent with good general combining ability for trait also exhibits good per se performance. This is true with parents S-22 and Local type for traits yield per plant, average fruit weight, number of locules per fruit, reducing sugar, total sugar, juice recovery percentage and plant height, primary branches number of flowers and fruits per truss, number of fruits per

plant, TSS, shape index, pulp content, reducing sugars respectively. Similar results were reported by Sonone et al. (1986) and Dhaliwal et al. (1999), Bhatt et al. (2001) Sharma et al. (2002) and Cheema et al. (2003). 5.2.2.3 Specific combining ability (sca) effects Plant height and number of primary branches are the important growth parameters which act as source trait to support yield and its component traits. The estimates of sca effects for two crosses for these two traits revealed that the crosses L-15 x PKM-1 (25.64) for plant height, Local x PKM-1 (1.34) for number of primary branches had significant and higher magnitude of sca effect. The Local type which is one of the parents in the cross Local x PKM1 was also a good combiner for number of primary branches and the cross involves positive x negative general combiners and the cross L-15 x PKM-1 had low positive x negative general combiners. Per se performance of the hybrid L-15 x PKM-1 is also highest. This may be due to intra allelic interaction (Niyaria and Bhalala, 2001). The next best crosses with desirable high sca effects for these traits were CO-3 x Solan Vajra for plant height and L-15 x PKM-1 for primary branches. In the cross CO-3 x Solan Vajra, the Solan Vajra is a good general combiner and CO-3 is not, where as in the cross L-15 x PKM-1, both are not good general combiners. Similar results reported by Mishra and Khanna (1977), Bhatt et al. (2001) and Cheema et al. (2003). The high total yield per plant is the ultimate motto of the breeder. In the present study, the top three crosses with high per se performance (S-22 x L-15, S-22 x Sivap and CO-3 x Solan Vajra) also exhibited high sca effects for total yield per plant. The cross S-22 x L-15 exhibited significant sca effect for number of flowers per truss, average fruit weight, total acidity reducing sugars, total sugars, juice recovery percent and pulp content. The parent S22 is a second top general combiner for the yield. S-22 x L-15 involved positive x positive general combiners, S-22 x Sivap involved positive x positive general combines and CO-3 x Solan Vajra involved negative x positive general combines for yield. S-22 parent having highest and significant gca effect, demonstrated its value as good general combiner for the total yield per plant. Further these two crosses viz., S-22 x L-15 and S-22 x Sivap having positive x positive gca effects reveals that the high gca effects in these crosses was mainly through additive gene effects. Therefore, the best option for improvement is the identification of transgressive segregants based on sca effects which may lead to isolation of promising lines of high total yield in tomato. Similar results were obtained by Sonone et al. (1986) and Cheema et al. (2003). The other cross CO-3 x Solan Vajra had revealed significant positive sca effect for plant height, number of fruits per plant and average fruit weight. This cross involved negative and positive combiners as its parents. In this cross genetic interaction might be additive x dominance. Good quality should be coupled with the yield for fetching better market price. The hybrid S-22 x Solan Vajra showed high positive and significant sca for pericarp thickness and also significant positive sca effect for traits TSS and juice recovery percentage. The hybrid, CO-3 x Local showed high positive and significant for TSS, and also exhibited significant positive sca effect for fruit shape index, reducing sugars and total sugars. One of the identified heterotic hybrid for yield over commercial check, CO-3 x Solan Vajra showed significant high and positive sca effect for ascorbic acid. The parent CO-3 showed high gca for fruit shape index, ascorbic acid where as Local type showed significant gca effect for traits like fruit shape index, ascorbic acid, reducing and non reducing sugar. The cross CO-3 x Local involves negative x positive gca effects for TSS hence gene action will be dominance x additive, for percarp thickness it is high negative x negative gca effects so non-additive gene effect is predominant here. The high gca effects in CO-3 x Solan Vajra cross was mainly due to additive gene effects. In general wherever Local type is involved as one of the parents, such crosses recorded significant and positive heterosis over mid parent, better parent commercial check for the characters viz., plant height, primary branches, flowers per truss, fruits per truss, fruits per plant. The involvement of Local type as one of the parents resulted in reduced average fruit weight and fruit yield (Plate 6). For the quality traits viz., TSS and ascorbic acid the involvement of Local type in the hybridization gave encouraging results as far as heterosis over mid parent, better parent and commercial check is concerned.

Local type is a good general combiner for traits like plant height, primary branches, number of fruits per plant, number of fruits per truss and number of flowers per truss. It is also a good general combiner for some of the quality traits like ascorbic acid, total acidity, TSS, pulp content giving scope for its utility in hybridization for developing hybrids for processing purpose. In general Solan Vajra is a good general combiner for traits like ascorbic acid and juice content indicating its use in producing hybrids that can enter into fresh market.

5.2.3 Characterization of the best heterotic hybrids for productivity

The crosses S-22 x L-15, S-22x Sivap and CO-3 x Solan Vajra were identified as the best heterotic cross combinations over commercial check. They showed significant positive standard heterosis ranging from 20.92 to 37.02. The relative performance of these hybrids in respect of 18 traits studied along with the check, MHTM-256 and the parental lines of these hybrids is given in Table 35. The three productive hybrids had significant higher per se value than the commercial check in respect of number of fruits per plant, average fruit weight, primary branches, pericarp thickness total acidity and juice content indicating that the higher productivity in these hybrids is associated with higher number of primary branches, number of fruits per plant and average fruit weight. The keeping quality is an important trait for perishable vegetable like tomato. The keeping quality increases with higher pericarp thickness of the fruits. The above identified hybrids exhibited higher pericarp thickness compared to commercial check. From processing point of view reducing sugars are important in tomato. The identified heterotic hybrid S-22 x L15 showed higher per se value over commercial check and the hybrid S-22 x Sivap showed per se value on par with that of commercial check. Another quality trait ascorbic acid is also improved in the crosses S-22 x Sivap and CO-3 x Solavajr as the per se values are higher than commercial check. Thus these hybrids are better than the commercial hybrid with respect to both productivity and quality. The table also reveals that the population mean for the characters number of fruits per truss, number of fruits per plant and TSS is higher compared to identified heterotic hybrids. This is because of the involvement of Local type genotype as it has smaller fruits, high number of fruits per plant and fruits per truss. Local type also showed higher TSS and the crosses involving Local type as one of the parents also showed similar results. But it is noteworthy that average fruit weight and yield was higher for identified heterotic hybrids compared to population mean per se value. The cross S-22 x L-15 also showed higher per se values for pulp content, total sugar and reducing sugar compared to population mean.

FUTURE LINE OF WORK

1. The superior segregants isolated through different mating schemes would serve as a better source material for further evaluation and finally to identify potential true breeding lines 2. Another cycle of intermating among the segregants with relatively high fruit weight and higher number of primary branches could also prove useful 3. The genotypes S-22, L-15, Sivap and Solan Vajra can be recommended for use as one of the parents to generate high yielding and better quality hybrids, as well as in varietal improvement programmes as they have high gca effect 4. It is suggested to test the superior hybrids along with few more in multilocation trial to confirm their potentiality and to know their stability over different agro-climatic situations 5. Selected parents with desirable per se and combining ability effects in respect to different component traits can be involved in multiple crossing scheme to recombine different productivity components

Table 35. Relative performance of top 3 hybrids with respect to per se value and check for 18 quantitative and quality traits in tomato

S-22 X L-15 Character S-22 Plant height (cm) Number of primary branches Number of flower/truss Number of fruits/truss Number of fruits/plant Average fruit weight (g) Number of locules/fruit Fruit shape index Pericarp thickness (mm) Ascorbic acid (mg/100g) Total acidity (%) Reducing sugar (%) Non-reducing sugar (%) Total Sugars (%) Juice recovery (%) Pulp content (%) Total soluble solids (° Brix) Yield per plant (g) 77.50 4.83 3.60 2.77 34.67 29.33 4.77 0.77 3.40 20.73 0.27 2.82 0.16 2.98 38.18 58.10 3.47 1010.00 L-15 93.30 4.92 3.33 2.60 32.67 23.20 2.55 0.95 3.90 21.05 0.36 2.79 0.21 3.00 30.30 67.00 4.30 763.33 F1 79.17 4.17 4.56 3.17 41.33 32.42 3.61 0.86 4.80 17.08 0.39 2.95 0.19 3.16 25.43 71.73 4.50 1333.33 S-22 77.50 4.83 36.50 2.77 34.67 29.33 4.77 0.77 3.40 20.73 0.27 2.82 0.16 2.98 38.18 58.10 3.47 1010.00 Sivap 89.73 4.75 3.27 2.55 33.33 29.47 3.42 0.74 3.60 16.92 0.34 2.54 0.57 2.83 34.40 61.23 4.48 983.00 F1 89.07 3.92 3.50 2.47 40.67 31.73 4.37 0.76 3.80 21.83 0.35 2.34 0.42 2.72 34.77 63.67 4.00 1288.70 CO-1 60.67 4.67 4.43 2.78 28.67 21.70 3.50 0.91 4.20 25.58 0.43 2.23 0.57 2.83 27.92 24.66 4.20 23.33 S-22 X Sivap CO-3 x Solan Vajra Solan Vajra 95.17 6.22 3.33 2.60 33.00 34.27 3.33 0.88 4.40 22.60 0.40 2.64 0.28 2.93 34.98 66.87 4.63 1131.67 F1 100.92 5.07 3.29 2.77 34.67 34.07 3.39 0.89 4.60 31.42 0.27 2.48 0.46 2.70 28.30 63.57 4.45 1176.67

Population Mean 88.48 4.96 4.24 3.23 49.40 23.90 3.26 0.83 4.85 23.45 0.35 2.48 0.36 2.86 31.40 66.90 4.85 910.00

MHTM-256

S.Em±

73.30 3.66 3.34 2.73 360.66 31.80 3.11 0.91 3.40 21.20 0.26 2.48 0.44 2.94 22.70 75.70 4.56 973.30

3.8 0.26 0.22 0.29 1.98 0.46 0.42 0.08 0.16 0.86 0.08 0.08 0.09 0.12 1.8 1.66 0.14 4.2

40 35 30 25 20 15 10 5 0

Percentage

S-22 X L-15

S-22 X Sivap Cross

CO-3 x Solan Vajra

Fig. 3. Per cent heterosis for top three hybrids for yield over commercial check

Fig. 3. Per cent heterosis for top three hybrids for yield over commercial check

Plate 5. Fruit features of top hterotic hybrids

Plate 6. Fruit features of hybrids involving Local type as one of the parents

VI. SUMMARY

BIPARENTAL MATING

The present investigation was carried out during the kharif season of 2005 at botany garden of Agricultural College, Dharwad to compare the relative efficiency of inter population mating over intra population mating and BIPs over F3 interms of release of genetic variability, frequency of superior segrgants and to know the shift in association pattern of component traits with fruit yield in tomato. In view of the above objectives, inter mating populations (within F2 of MHTM 256 and S-4-14) (population M and population S). Inter mating populations (using MHTM 256 as female and S-4-14 as male and vice versa crossed in F2) (population M/S and S/M) and F3 were evaluated. The experiment was laid out without replication by transplanting the seedlings with spacing of 75 cm between rows and 60 cm between plants within row. 1. The comparison of mean of characters among segregating populations indicated that mean values in general were relatively higher in case of inter mating BIP population compared to intra mating population and the mean values of BIPs were relatively higher than F3. Among BIPs, population S showed higher mean for most of the traits. As regard to the range of expression for the traits is concerned wide range was noticed in inter mating population compared to intra mating population and BIPs showed wide range than selfed progeny. It is noteworthy that especially the upper limit of range was higher in magnitude for all characters. At the same time the lower limit was smaller compared to that of commercial check and F3 population, suggesting the role of inter mating in releasing more variability than in selfed generation. Among BIPs, population M/S showed wider range for traits like fruits per plant and fruits yield per plant. Though intra mating population showed high GCV and PCV for traits like primary branches, fruits per truss and pericarp thickness but for the most important traits like yield per plant and average fruit weight the inter mating population showed high GCV and PCV. The BIP showed high GCV and PCV for all the characters except fruits per plant, yield per plant and fruit shape index compared to F3. Among the characters, fruits per plant and primary branches showed high GCV and PCV indicating presence of repulsion phase linkage for these traits. Among the populations population M/S (inter mating population) showed higher GCV and PCV for most of the traits under study. The heritability estimates and genetic advance as per cent of mean also showed similar results as that of GCV and PCV. Among the characters average fruit weight showed highest heritability estimate and genetic advance as per cent of mean followed by yield per plant, fruits per plant, plant height and primary branches. Among the BIP populations, population M and population M/S showed high heritability and GA as per cent of mean for important yield and yield attributing traits. Analysis of correlation co-efficient revealed high positive association of fruits per plant with fruit yield irrespective of type of population. Primary branches had higher association next to fruits per plant. Interestingly, inter mating resulted in the shift in correlation from non-significant to significant association. Path analysis revealed that fruits per plant showed maximum direct effect on fruits per plant in all the populations. The number of primary branches indirectly affected yield via number of fruits per plant, thus indicating that, fruits per plant is one of the most important yield contributing trait in tomato. Hence, in any breeding programme, selection based on number of fruits per plant and high average fruit weight may be expected to increase the fruit yield. In general, inter mating population showed higher frequency of superior segregants than intra mating population for yield per plant and number of superior segregants were higher in population S/M. When compared to F3, BIPs showed higher number of superior segregants for the character stated above. The study clearly highlighted the distinct advantage of inter mating over intra mating and biparental mating over selfing series in a self pollinated species like tomato.

2.

3.

4.

5. 6.

7.

8.

9.

10. The comparison of biparental mating and selfing generation and of intermating and intra mating BIP population shows that whatever additional variability generated through the use of inter mating in the early segregating generations has been the consequence of release of concealed variability, in the segregating generations, which is probably brought about by rare combination between the tightly linked genes. Inter mating was also found to be effective in changing not only the magnitude and direction of correlation coefficient but also direct and indirect dependence of fruit yield on its component traits. This approach will help in creating new population with high frequencies of rare recombinants which can not otherwise be realized in small segregating populations normally being raised through the conventional method of breeding especially when desired genes are unfavourably linked. In addition to these, it will also help in maintaining a greater variability for selection to be effective for longer period and will thus avoid the early fixation of genes in homogenous state.

DIALLEL MATING

A field experiment was under taken to study heterosis and combining ability in tomato (Lycopersicon esculentum Mill.) with the objectives of identifying good general combiners and to asses the magnitude of heterosis for yield and yield related components. The experiment was conducted in Botany Garden, Department of Genetics and Plant Breeding, University of Agricultural Sciences, Dharwad. The salient features of the experimental findings are summarized below. Twenty-one hybrids were developed by crossing seven parents following half diallel mating without reciprocals. All the hybrids were evaluated along with the parents in Randomized Block Design (RBD) with three replications. Variance due to parents and hybrids were significant for all the yield and yield related components. The magnitude of heterosis over commercial check (MHTM-256) was very high. It is ranged from -29.69 to 61.43 per cent for plant height, -5.73 to 91.25 per cent for primary branches, -2.69 to 67.3 per for flowers per truss; 2.47 to 4.99 per cent for fruits per truss, 18.46 to 308.77 per cent for fruits per plant, -78.89 to 18.26 per cent for average fruit weight; 33.88 to 37.02 per cent for fruit yield per plant. The magnitude of heterosis of quality parameters like TSS, ascorbic acid content, total acidity, reducing sugars, non-reducing sugars, total sugars, juice content and pulp content ranged from -12.2 to 39.7 per cent, 14.50 to 31.70 per cent, 3.84 to 69.2 per cent, 10.5 to 18.95 per cent, -61.3 to 29.54 per cent, -11.22 to 9.67 per cent, 19.25 to 79.18 per cent and -23.56 to -6.58 per cent, respectively. For the characters locules per fruit, fruit shape index and pericarp thickness the magnitude of heterosis ranged from -27.33 to 47.44 per cent, -20.8 to 12.08 per cent and -35.29 to 35.29 per cent respectively. Out of 21 hybrids, seven, three and three hybrids exhibited positively significant heterosis for total yield per plant over mid parent, better parent and standards check hybrid respectively. Maximum standard heterosis for total yield per plant was observed in the cross S-22 x L-15 (37.02%) followed by S-22 x Sivap (32.44%) and CO-3 x Solan Vajra (20.92). The S-22 x L-15 also exhibited significant standard heterosis for growth parameters and yield components. Among the seven parents evaluated, the highest and significant gca effect for yield per plant was observed in S-22 followed by Sivap, L-15 and Solan Vajra. S-22 and Sivap also exhibited maximum gca effect for average fruit weight, reducing sugars, non-reducing sugars total sugars and juice content. Though Local type exhibited negative gca effect for yield per plant, it showed positive and significant gca effects for plant height, number of primary branches, number flower and fruits per truss, number of fruits per plant, TSS, shape index, pulp content and reducing sugars. The hybrids L-15 x PKM-1 and CO-3 x Solan Vajra were identified as good specific combiners for plant height. Similarly Local x PKM-1 and L-15 x PKM-1 for primary branches, S-22 x L-15 and L-15 x Solan Vajra for number of flowers per truss, Sivap x Local and Local x PKM-1 for primary branches, S-22 x L-15 and L-15 x Solan Vajra for number of flowers per truss, Local x PKM-1 and S-22 x Local for fruits per plant L-15 x PKM-1 and CO-3 x Solan Vajra for average fruit weight and CO-3 x Solan Vajra, S-22 x L-15 and S-22 x Sivap were identified as best specific combiners for yield per plant. CO-3 x Local and S-22 x Solan Vajra

for TSS, Sivap x PKM-1 for locules per fruit, Solan Vajra x PKM-1 and Sivap x Local for fruit shape index, S-22 x Solan Vajra for pericarp thickness CO-3 x Sivap and CO-3 x Solan Vajra for ascorbic acid, S-22 x PKM-1, L-15 x Solan Vajra and CO-3 x Sivap for total acidity were good specific combiners. For reducing sugar CO-3 x PKM-1 and CO-3 x Local identified as specific combiners. Similarly L-15 x Local and CO-3 x Sivap for non-reducing sugars, S-22 x L-15 and CO-3 x Local for total sugar, L-15 x Local and S-22 x L-15 for juice content and L-15 x Sivap and Local x PKM-1 for pulp content are good specific combiners. Evaluation of parents across all the traits studied indicated Local type, S-22, Solan Vajra and L-15 as good general combiners. Similarly evaluation of hybrids over all the 18 traits revealed S-22 x L-15, L-15 x CO-3, CO-3 x Sivap and CO-3 x Solan Vajra are the best specific combinations.

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GENETIC AND BREEDING INVESTIGATIONS IN TOMATO (Lycopersicon esculentum Mill.)

PURNANAND G. KULKARNI 2006 ABSTRACT

Tomato (Lycopersicon esculentum Mill.), an extremely popular vegetable around the world is the focus of large agricultural industry. Two investigations viz., biparental mating and diallel mating were carried out during kharif 2005-06 in Botany garden of Agriculture College, Dharwad. Biparental mating was carried out to assess the variability generated and to compare the efficiency of inter population biparental mating over intra population biparental mating and of biparental mating over selfed population for yield and its important components. Intrapopulation mating involved the crosses within each F2 population of MHTM-256 and S-414 while interpopulation mating was carried out between selected F2 plants of MHTM-256 as male with selected F2 plants of S-4-14 and vice-versa. Of the two the best F2 population (S4-14) was selfed to get F3 progeny. The results revealed that mean values, in general were high in F3 than BIPs because of wider variability generated in BIPs especially intermating populations. The magnitudes of GCV, PCV, heritability and genetic advance were enhanced in BIPs for all the characters studied than the selfed population. Among BIPs, inter population mating exhibited high heritability and genetic advance for plant height, average fruit weight and yield. Biparental mating also resulted in shift in the magnitude as well as direction of correlation coefficients. Diallel mating was carriedout with seven parents and 18 quantitative and quality traits were evaluated. The results indicated that, heterosis over midparent, better parent and commercial check was exhibited by all the characters. S22 x L-15, S-22 x Sivap and Co-3 x Solan Vajra were the top heterotic hybrids for yield over check (MHTM-356). Combining ability analysis revealed that, S-22, Solan Vajra, Local type and L-15 were good general combiners whereas S-22 x L-15, S-22 x Sivap and Co-3 x Solan Vajra were the best specific combiners.

Dr. O. SRIDEVI MAJOR ADVISOR

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