that flowering starts annually in the same calendar period, irrespective of ..... reported a wide range of variability at both fertility levels in twenty-eight true breeding ..... violet x white crosses showed violet flowers in the F1 progeny, and the ratio ...
DEDICATION I DEDICATE THIS HUMBLE EFFORT, THE FRUIT OF MY THOUGHTS AND STUDY TO MY YOUNGER SISTER (LATE)
ASSESSMENT OF GENETIC VARIATION AND INHERITANCE OF YIELD AND YIELD RELATED TRAITS IN LENTIL (Lens culinaris Medik.)
A THESIS SUBMITTED TO THE UNIVERSITY OF THE PUNJAB IN FULFILMENT OF THE REQUIREMENT FOR THE DEGREE OF DOCTOR OF PHILOSOPHY (Ph. D) IN BOTANY
By MUHAMMAD ASHRAF
Department of Botany, University of the Punjab, Lahore, Pakistan 2008
TABLE OF CONTENTS Chapter
Title
Page i
Acknowledgements Abstract List of Tables List of Figures
iv
1
Introduction
1
2
Materials and Methods
34
2.1 Assessment of Genetic Variation in Lentil Germplasm
35
2.2 Estimation of Genetic Diversity in Parental Lines
39
ii
vi
2.2.1
Estimation of genetic diversity in parental lines using morphological characteristics
39
2.2.2
Estimation of genetic diversity in parental lines using randomly amplified polymorphic DNA (RAPD) markers
40
2.3 Estimation of Nature and Pattern of Inheritance of Quantitative and Qualitative Traits
43
2.3.1
Hybridization for production of F 1 , F 2 , BC 1 and BC 2 populations
43
2.3.2
Raising populations for inheritance studies
44
2.3.3
Data recording for estimation of nature of inheritance on quantitative traits and pattern of inheritance on qualitative traits, and statistical analyses
45
Chapter
Title
Page
3
Genetic Variation in Lentil Germplasm
47
4
Genetic Diversity in Parental Lines
75
5
Nature of Inheritance for Different Plant and Yield Related Traits
91
6
Pattern of Inheritance for Qualitative Traits
155
7
Discussion and Conclusions
166
References
181
Appendices
209
Acknowledgements Above everything else, I exalt and bow my head before “ALLAH” The most beneficent and merciful. I offer my humblest thanks to Him for bestowing upon me. His blessings with intellectual capability to work hard and successful accomplishment of my dissertation. Trembling lips and wet eyes praise for Holy Prophet Muhammad (PBUH) who is forever beacon of guidance and knowledge for humanity as a whole and enables us to recognize our creator. The author feels pleasure and honour of this available opportunity to offer sincerest thanks and gratitude to his worthy supervisor Professor Dr. Shahida Hasnain, Dean Faculty of Life Sciences, University of the Punjab, Lahore for her kind and affectionate guidance and enlightened supervision during my research work. Her sympathetic and pleasant attitude has always been a consistent source of inspiration. I offer thanks to my co-supervisor, Dr. Muhammad Siddique Sadiq, Deputy Chief Scientist (Rtd), NIAB, Faisalabad for his kind cooperation and guidance. I offer my cordial and profound thanks to Dr. Ilyas A. Malik, PSO (Rtd), NIAB, Faisalabad for his kind guidance and teaching attitude. I offer my cordial and profound thanks to Mr. Akbar Ali Cheema, Deputy Chief Scientist, NIAB, Faisalabad for his help and providing facilities. I feel pleasure in expressing my sincere thanks to Mr. Ghulam Rasul Tahir, Deputy Chief Scientist (Rtd) and Mr. Muhammad Akram, ARO, NIAB for their help in statistical analysis. I also thankful to Mr. M. D. Idrees, Sr. Librarian, NIBGE, Faisalabad for his help in literature search. I feel pleasure to thank Dr. W. Erskine (now Director, CLIMA, Australia) and Dr. A. Sarker, International Center for Agricultural Research in the Dry Areas (ICARDA), Syria for their kind help and cooperation. Last, but not the least, I shall no be loyal to my family, if I do not admit and mention the source of my real pleasure which enlightened my mind and spirit. I offer humble gratitude to my loving father, mother, brothers, spouse and kids whose hearts beats with golden sentiments and whose hands always arise in pray for my success.
MUHAMMAD ASHRAF i
Abstract Lentil germplasm from Pakistan is of the pilosae group with marked lack of genetic variability and paucity in variation within this germplasm came from the sensitivity of a world lentil collection to temperature and photoperiod. In comparison with other lentils, this group is characterized by morphological traits, precocity in flowering and maturity, low biomass and increased sensitivity of the temperature. The asynchrony in flowering between exotic material and the indigenous germplasm enforced reproductive isolation, and this bottleneck has played a major role in reducing lentil seed yield in the region. Therefore, understandings of the genetic relationships and diversity of lentil in relation to germplasm from other countries to identify important agronomic traits, estimation of the components of genetic variation for different plant/yield related characteristics in the segregating populations, and determination of the type of gene action for important qualitative/quantitative traits for the selection of desirable recombinations with stable and improved seed yield are the important aspects that might play pivotal role in improving the productivity and stability of the lentils. Knowledge about the germplasm diversity and genetic relationships among breeding materials may be an invaluable aid in improvement strategies. The present studies, comprising evaluation of germplasm to estimate the genetic variation for different plant and yield contributing traits, evaluation of parental lines on both morphological and molecular markers, and their use in the production of segregating populations for inheritance studies of different quantitative as well as qualitative key characters, were carried out for their implications in the future breeding strategies. Data of lentil germplasm and parental lines were subjected to various biometrical approaches to determine the key factors/traits contributing to improved seed yield under the prevailing growth conditions, and the diversity among the parental lines. Highly significant differences observed among the genotypes for the characters under study indicated that the germplasm was amenable to improvements. In the present studies, phenology was observed as a key plant trait influencing seed yield as high temperatures during pod/seed development phase(s) through forced maturity cause low pod setting and low seed weight resulting in poor seed yield. Positive associations of seed yield per plant with fertile nodes on primary and secondary branches, seeds per pod, pods per plant, 100 seed weight, biomass per plant, and harvest index in relation with direct effects of biomass followed by pods per plant, fertile nodes on secondary branch, 100 seed weight and secondary branches
ii
per plant indicated the importance of these characters during selection. Whereas, negative correlations of days to flower with fertile nodes on primary and secondary branches, pods per plant, 100 seed weight, biomass and seed yield per plant revealed that selection based on seed yield contributing traits alone may not be fruitful if the phenology of the plants do not synchronize with the prevailing growing climate. The negative association of days to flower with seed yield per plant was also confirmed by the negative direct effect of harvest index on seed yield per plant. The associations of yield related traits (pods per plant, seed weight, pods per plant, 100 seed weight, biomass per plant and harvest index) and days to flower with seed yield per plant showed the importance of these traits that should receive due consideration while selecting lentil genotypes for higher seed yield. Bushy as well as determinate plant types may contribute towards selection for improved seed yield. However, the expression of productivity under field conditions and the translation of this improvement to higher seed yields depend on the environment and phenology, and the canopy structure. The parental lines used for the production of segregating populations for inheritance studies generally showed relatively low genetic diversity. Likewise, nonlinear relationships among the parents based on their geographic pattern and genetic diversity as well as with their visible characters were also observed. Genetic analysis using generation means revealed significant difference among generations. Dominance gene effect (h) was found to be more prominent than the additive gene effect (d) in most crosses. The epistasis interactions (i, j and l) were observed prominent in controlling the inheritance of different plant traits. Seed yield per plant and its contributing traits were controlled by both additive and non-additive interactions (additive, dominance, and epistasis) suggesting that appropriate selection procedures may be adopted to improve the traits according to their mode of inheritance and the materials. Presence of non-additive gene interactions in most of the plant characters as well as seed yield per plant and its contributing components indicated that conventional selection procedure may not be effective enough for improvement of seed yield. Therefore postponement of selection in later generations or intermating among the selected segregants followed by one or two generations of selfing could be suggested to break the undesirable linkage and allow the accumulation of favorable alleles for the improvement of this trait. An improved knowledge of the genetic basis of qualitative characters aimed to select new cultivars carrying appealing traits will better support the lentil breeding programmes. The integration of classical plant breeding with molecular markers and QTLs mapping for genetic studies of different traits of interest also seemed beneficial to assist in the lentil improvement.
iii
LIST OF TABLES Table
Title
Page
2.1
Details of lentil germplasm evaluated at the Nuclear Institute for Agriculture and Biology (NIAB), Faisalabad, Pakistan, during 200203
36
2.2
Meteorological data for the growing season of lentil germplesm during 2002-03
37
3.1
Values of mean square, range, means and coefficient of variance for different plant characteristics in lentil germplasm
48
3.2
Genotypic (G) and phenotypic (P) correlations (r) among various plant characteristics in lentil germplasm
50
3.3
Direct and indirect effects of different characteristics on seed yield in lentil germplasm
53
3.4
Multiple correlation analysis of seed yield with different combinations of characters in lentil germplasm
56
3.5
Estimates of phenotypic and genotypic variances, heritability and genetic advance for different plant characters in lentil germplasm
58
3.6
Principal components analysis (PCA), varimax rotated factor loadings and communalities for variation among sixteen quantitative traits in lentil germplasm
60
4.1
Values of means, coefficient of variance, mean square, and range for different plant characteristics in lentil parental lines
77
4.2
Principal components (PCA) analysis, varimax rotated factor loadings and communalities for eleven quantitative traits in lentil parental lines
79
4.3
Estimates of similarity matrices in the parental lines using RAPD markers
84
5.1
Mean square values for different plant characteristics in 12 crosses of lentil (Lens culinaris Medik.)
93
5.2
Estimates of additive, dominance and non-allelic gene effects for days to flower in 12 lentil crosses
95
5.3
Estimates of additive, dominance and non-allelic gene effects for days to mature in 12 lentil crosses Estimates of additive, dominance and non-allelic gene effects for plant height in 12 lentil crosses
98
5.4 5.5
Estimates of additive, dominance and non-allelic gene effects for number of primary branches per plant in 12 lentil crosses iv
101 104
Table
Title
Page
5.6
Estimates of additive, dominance and non-allelic gene effects for number of secondary branches per plant in 12 lentil crosses
108
5.7
Estimates of additive, dominance and non-allelic gene effects for number of nodes on primary branch in 12 lentil crosses
111
5.8
Estimates of additive, dominance and non-allelic gene effects for number of fertile nodes on primary branch in 12 lentil crosses
114
5.9
Estimates of additive, dominance and non-allelic gene effects for number of nodes on secondary branch in 12 lentil crosses
117
5.10
Estimates of additive, dominance and non-allelic gene effects for number of fertile nodes on secondary branch in 12 lentil crosses
121
5.11
Estimates of additive, dominance and non-allelic gene effects for pod length in 12 lentil crosses
124
5.12
Estimates of additive, dominance and non-allelic gene effects for pod breadth in 12 lentil crosses
127
5.13
Estimates of additive, dominance and non-allelic gene effects for seeds per pod in 12 lentil crosses
129
5.14
Estimates of additive, dominance and non-allelic gene effects for pods per plant in 12 lentil crosses
132
5.15
Estimates of additive, dominance and non-allelic gene effects for 100 seed weight in 12 lentil crosses
135
5.16
Estimates of additive, dominance and non-allelic gene effects for biomass per plant in 12 lentil crosses
138
5.17
Estimates of additive, dominance and non-allelic gene effects for harvest index in 12 lentil crosses
141
5.18
Estimates of additive, dominance and non-allelic gene effects for yield per plant in 12 lentil crosses
144
6.1
157
6.2
Segregation pattern and P values for seedling growth habit and flower initiation in lentil Segregation pattern and P values for hypocotyls colour lentil
6.3
Segregation pattern and P values for seed coat pattern in lentil
159
6.4
Segregation pattern and P values for cotyledon colour in lentil crosses
161
7.1
Assessment of genetic variation through analysis of variance, correlations, path analysis and genetic parameters for important plant characteristics in lentil germplasm
168
v
158
LIST OF FIGURES Figure
Title
Fig 4.1
Hierarchical clustering among morphological plant traits
on
81
Fig 4.2a
Amplification of RAPD marker OPN-06 for lentil parental lines (1. ILL 4605, 2. ILL 5782, 3. ILL 6024, 4. Pant L 406, 5 ILL 8117, 6. ILL 7556, 7. ILL 7715, 8. ILL 6468, 9. ILL2580, 10. Masoor 93, 11. Turk Masoor, 12. ILL 6821)
83
Fig 4.2b
Amplification of RAPD marker OPB-1 for lentil parental lines (1. ILL 4605, 2. ILL 5782, 3. ILL 6024, 4. Pant L 406, 5 ILL 8117, 6. ILL 7556, 7. ILL 7715, 8. ILL 6468, 9. ILL2580, 10. Masoor 93, 11. Turk Masoor, 12. ILL 6821)
83
Fig 4.2c
Amplification of RAPD marker OPA-13 for lentil parental lines (1. LL 4605, 2. ILL5782, 3. ILL 6024, 4. Pant L 406, 5 ILL 8117, 6. ILL 7556, 7. ILL 7715, 8. ILL 6468, 9. ILL2580, 10. Masoor 93, 11. Turk Masoor, 12. ILL 6821)
83
Fig 4.3
Dendrogram showing genetic relationships among lentil parentals based on RAPD markers
85
vi
lentil
Page parentals
based
Chapter 1
INTRODUCTION Lentil (Lens culinaris Medik.) is an important grain legume crop which is cultivated worldwide as human food providing an important source of dietary proteins (Muehlbauer et al. 1995). Recent developments in dietary fashions favouring high-fibre foods and plant foods rich in protein can be expected to increase demand for lentils (Smart, 1990). The optimal climate for the crop is basically temperate. In Pakistan, the latter part of the reproductive growth phase in lentil often coincides with increasing temperature and dry conditions in Pakistan. This exaggerates the indeterminate nature of the crop, leading to immature pod/seed development, and subsequently resulting in forced maturity with low seed yield. The length of the growing season in Pakistan is delimited between the onset of winter (late October/early November) and spring (mid April), such that no significant window exists to extend the length of the growing period to meet the requirement of a long growing season in this crop. Early crop maturity is essential to match crop duration with the period of favorable growing conditions, avoid losses caused by increasing temperature, and to stabilize yield and quality. South Asia grows almost half the world's lentils, but the productivity of the crop in the region has historically been poor mainly due to inherent poor yield potential, loss of genes for higher productivity, and susceptibility to biotic and abiotic stresses (Muehlbauer et al. 2006; Gupta and Sharma, 2006), that breeders need to work on to improve yields. Moreover, differences in the length of the growing season have always made it hard to incorporate genetic material from elsewhere (Erskine and Manners, 1994). Selection on the basis of seed yield (a polygenic complex character) is usually not very efficient, but 1
selection based on its component characters could be more efficient. Plant breeders always focused their attention to the genetic improvement of the lentil plant both at national and international level. As a result, endeavours to improve lentil have been promising in some cases (Erskine, 1983), but this was not the case everywhere (Dutta and Mondal, 1998). Great effects of edaphic and climatic conditions on different morphological as well as yield contributing characters in lentil have also been observed (El-Gawad et al. 1997; Manara and Manara, 1983; Singh and Saxena, 1982). During migration of a landrace, especially towards isolated areas, reduction of genetic diversity often occurred as a result of bottleneck effect (Sonnate et al. 1994; Laghetti et al. 2005). Direct introduction into Indo-Pak subcontinent suffered from the problems of morphophysiological and biochemical characteristics with West Asian germplasm. The asynchrony in flowering has isolated the local pilosae ecotype reproductively. This was partly broken by the arrival of the early flowering cultivar ‘Precoz’ (Erskine and Saxena, 1993; Erskine et al. 1998) but the interaction of this cultivar (belonging to Argentina) with the principal climatic elements (temperature and photoperiod) has been observed to have significant inverse association with days of birth to flowering suggesting it sensitive to high temperatures (Gray and de Delgado, 1986).
Origin, Taxonomy and Cytology The putative progenitor of the cultivated lentil is Lens culinaris subsp. orientalis (Boiss.) Ponert which was distributed from Greece in the west to Uzbekistan in the east, and from the Crimean Peninsula in the north to Jordan in the south (Ladizinsky, 1979a; Cubero, 1981). The oldest carbonized remains of lentil were from Franchthi cave in Greece dated to 11,000 BC and from Tell Mureybit in Syria dated 8500-7500 BC (van Zeist in Zohary, 1992; Hansen and Renfrew, 1978). But as it is not possible to differentiate wild from cultivated small-seeded lentil, the state of domestication of these
2
and other carbonized remains in the aceramic farming villages in the 7th millennium BC in the Near East arc is unknown. The finding of a large hoard of lentil (about 1.4 million seeds) at Yiftah-el dated to 6800 BC was, however, suggestive of domestication (Garfinkel et al. in Zohary, 1992). The oldest find of lentil seeds that were larger than wild seeds, and therefore unequivocally domesticated (Helbaek, 1969), was at Tepe Sabz, Iran, and have been dated to 5500-5000 BC. The overlap in the distribution of wild lentil and the early archaeological record indicates that lentil was domesticated in the Near East arc (Erskine, 1998). Medikus is considered the authority for cultivated lentils because the publication of L. culinaris Medikus predates that of L. esculenta Moench (Slinkard, 1974). Cultivated lentil belongs to the genus Lens that is associated with other genera of the Vicieae tribe (Kupicha, 1981). The primary gene pool of Lens culinaris comprises ssp. culinaris and its presumed wild progenitor ssp. orientalis. Three other wild species are recognized in the secondary gene pool and include L. odemensis Ladizinsky, L. nigricans (M. Bieb.) Gorden and L. ervoides (Brign.) Grande (Ladizinsky, 1993). Ahmad and McNeil (1996) reported that Lens culinaris subsp. orientalis appeared as the wild progenitor of the cultivated lentils whereas Ahmad et al. (1997a) reported that L. culinaris subsp. orientalis and L. odemensis were probably the wild progenitors of cultivated lentil. Ferguson and Erskine (2001) and Ferguson et al. (2000) reported that the genus Lens comprised of seven taxa in four species and Lens orientalis is the presumed progenitor of cultivated L. culinaris and the two species are fully crossable and produce fully fertile progenies. Jeena and Singh (2001a) suggested that L. ervoides was separated from L. orientalis (L. culinaris subsp. orientalis) and L. odemensis (L. culinaris subsp. odemensis) on the first principal axis. L. orientalis and L. odemensis were close to each other and were separated from L. nigricans. On the first vs third principal axes, most of the L. orientalis accessions
3
fell on the periphery, while L. ervoides and L. nigricans fell near the centre. Work has triggered the recognition of two new additional species of Lens: L. lamottei Czefranova and L. tumentosus Ladizinsky (Ladizinsky, 1997a; van Oss et al. 1997). Studies of seed surface ornamentation and seed protein profiles in wild and cultivated lentil taxa (Mallick and Sawhney, 2002 & 2003) supported the view of L. culinaris subsp. orientalis being the probable wild progenitor of cultivated lentils. Mayer and Bagga (2002) reported that the basal and highly divergent status of the L. nigricans clade is depicted, and the progenitor-derivative relationship between L. culinaris subsp. orientalis and L. culinaris subsp. culinaris is reaffirmed. Phylogenetic analysis confirmed the divergence of L. nigricans from all species, and the closeness of cultivated lentil to its wild progenitor, although two gene pools could possibly be identified in subsp. orientalis (Sonnante et al. (2003). Barulina (1930) classified the assembled variation into six groups (grex varietatum), each of which was geographically differentiated and also characterized by a complex of morphological characters, mainly qualitative, common within a group but differing in other groups. Lentil landraces were classified into macrosperma and microsperma types according to seed size and an array of associated characters generally insensitive to the environment. Microsperma types are characteristic of the Indian subcontinent, parts of the Near East, and the lower latitudes of the Old World, including Ethiopia and Yemen. Geographic differentiation between landraces also has been found for other more cryptic, eco-physiological factors, such as those resulting from selection for soil conditions and for climatic conditions. All Lens species are self-pollinating annual diploids (2n = 14) (Barulina, 1930; Ladizinsky, 1979a; Ladizinsky et al. 1984). Lens species have similar karyotypes consisting of three pairs of metacentric or submetacentric chromosomes, three pairs of
4
acrocentric chromosomes and one satellite pair of chromosomes (Ladizinsky, 1979a; Sharma and Mukhopaday, 1963; Slinkard, 1985).
Domestication and Distribution Archaeological studies have confirmed the presence of lentil in the Turkey-SyriaIraq region as far back as 8500–600 BC. This region probably played an important role in lentil domestication and starting the further spread to the Nile, Greece, Central Europe and eastwards to South Asia (Nene, 2006). The wild lentil species survived in small disjunct populations, predominantly in the Mediterranean basin, although L. culinaris Medik. subsp. orientalis (Boiss.) Ponert stretches east to Turkmenistan. The geographical distributions and ecological preferences of the taxa were reviewed by Ladizinsky (1993) and also reported in the Eastern Mediterranean region as one of the first domesticated plants (Zohary and Hopf, 1988). The crop spread to the Nile, and to Central Europe via the Danube. The crop was part of the assemblage of Near Eastern grain crops introduced to Ethiopia by the invaders of the Hamites. From the Bronze Age onward, maintained itself as an important companion of wheat and barley throughout the expanding realm of Mediterranean-type agriculture. The dissemination eastward of the Near Eastern grain crops, including lentil, reached Georgia in the 5th and early 4th millennia BC. Lentil cultivation spread rapidly with that of Neolithic agriculture to the Nile Valley, Europe and Central Asia. After 1500 A. D. the Spanish introduced lentil to South America via Chile (Solh and Erskine, 1984). The crop probably reached its current Old World range about 3000 years ago. It was carried to the New World after Columbus. The crop appeared in the archaeological record in India around 2500 BC as part of the Harappan crop assemblage. Alphonse de Candolle (1882) wrote that on linguistic grounds, “It may be supposed that the lentil was not in this country (India) before the invasion of the Sanskritspeaking race”. Evidence of the spread of lentil eastward into the Indo–Gangetic Plain
5
dates to around 2000 B. C., but previous contacts between Mohenjo–Daro, and the Sumarians and Akkadians of Mesopotamia are well documented, and it may have been introduced into the Indus valley earlier (Cubero, 1981), where it was exposed to new, markedly different environmental conditions. Flowering responses to temperature and photoperiod of a world collection illustrated the responses to selection for adaptation to new ecological environments following the spread of the crop from its origin (Erskine et al. 1994a). Dissemination to lower latitudes such as into Egypt, Ethiopia and India was accompanied by a reduction in photoperiodic response. Obligate photoperiodic control of the onset of flowering ensures that flowering starts annually in the same calendar period, irrespective of fluctuations in temperature. Consequently, selection against photoperiodic control in a long-day plant such as lentil implies an adaptation to relatively short days, which occur at low latitudes and which would otherwise delay flowering to an unacceptable extent. Under these conditions, the crop relies rather more on temperature than photoperiod to ensure that flowering occurs at an ecologically and agronomically appropriate time. On average, there was also evidence that flowering in the subtropical group was more temperaturesensitive than in West Asian germplasm (Erskine, 1998). Temperature sensitivity to processes other than flowering, such as the base temperature for rate of germination (Tb), also has been altered during dissemination from the Near East to environments with higher temperature regimes. The mean Tb of 15 randomly selected accessions from Ethiopia and India was higher than that of similar samples from Lebanon and Turkey (Ellis and Hong, 1995). Regional differences resulted from the spread of the crop to new physical environments with the consequent natural and artificial selection for local adaptation. The spread of lentil from the Near East into India resulted in a loss of winter hardiness and the ability to extract iron from calcareous,
6
high pH soils, a reduction in photoperiodic sensitivity for flowering, increased intrinsic earliness in flowering, an increase in temperature sensitivity for flowering, and in the base temperature for rate of germination. Each factor individually might be of minor importance, but collectively they illustrated the complex of interacting ecophysiological factors that determined adaptation.
Morphology and Floral Biology The cultivated lentil (Lens culinaris Medik.) is an annual bushy herb, almost erect or semi-erect, much branched, softly hairy; stems slender, angular, and 15-75 cm height (Duke, 1981; Muehlbauer et al. 1985). Ten to sixteen leaflets are subtended on the rachis (40-50 mm); upper leaves have simple tendrils while lower leaves are mucronate (Muehlbauer et al. 1985). The leaves are alternate, compound, pinnate, usually ending in a tendril or bristly; leaflets 4-7 pairs, alternate or opposite; oval, sessile, 1-2 cm long; stipules small, entire; stipules absent; pods oblong, flattened or compressed, smooth, to 1.3 cm long, 1-2-seeded; seed biconvex, rounded, small, 4-8 mm × 2.2-3 mm, lens-shaped, green, greenish-brown or light red speckled with black; the weight of 100 seeds range from 2 to 8 g; cotyledons red, orange, yellow, or green, bleaching to yellow, often showing through the testa, influencing its apparent color (Kay, 1979; Duke, 1981; Muehlbauer et al. 1995). Flowering begins acropetally and the lowermost buds open first and flowering proceeds upward and it takes about two weeks to complete opening of all the flowers on the single branch (Nezamudhin, 1970). The opening of flower occurs between 8.00 to 10.00 hrs and continues till noon and each flower remains open for about 16–24 hrs. At the end of the second day and on the third day all the opened flowers close completely and the colour of the corolla begins to fade. The setting of pods occurs after 3–4 days. The flowers have small ovaries with one or two ovules. The style is covered with a hairy inner surface. Each flower produces a short pod containing one or two lens-shaped seeds. Flowers can be
7
white, lilac or pale blue in color and are self-pollinated. At maturity plants tend to lodge because of their weak stems (www.hort.purdue.edu/newcrop/afcm/lentil.html). Flowers are small, pale blue, purple, white or pink, in axillary 1-4-flowered racemes; 1-4 flowers are borne on a single peduncle and a single plant can produce up to 10-150 peduncles each being 2.5-5 cm long. The size of seeds increases from the types grown in eastern regions to western types. Two types, namely; macrosperma, found mainly in the Mediterranean region and the New World (seed size ranging from 6 to 9 mm in diameter and yellow cotyledons with little or no pigmentation), and microsperma (2 to 6 mm with red orange or yellow cotyledons) found on the Indian subcontinent, Near East and East Africa, respectively, are known (Muehlbauer et al. 1985). The first one includes the Chilean or yellow cotyledon types while the latter includes the small seeded Persian or red cotyledon lentils (Kay, 1979).
Studies on Genetic Variation/Diversity Grain yield is a complex phenomenon controlled by a number of morpho-genetic characters and is highly influenced by the environment. Quantitatively genetic parameters such as heritability and variance components are useful for designing new breeding programmes, predicting response to recurrent selection, allocating resources in field performance trials and constructing selection indices. Correlation coefficients between the features are useful because they give information about the effect of the selection on other traits whereas indirect selection can be advisable if this is more practicable, other features are influenced positively, the genetic correlation is narrow between the two traits if the correlated trait has a higher heritability than the trait therein selected. The selection success can be estimated in the correlated feature if the heritabilities of both traits and the genetic correlation between them are known (Falconer, 1984). Muehlbauer (1974) observed that seed weight was negatively correlated with yield whereas single, double,
8
triple, and quadruple-pods/plant, yield/plant, seeds/plant, pods/plant, and seeds/pod were positively correlated suggesting that recurrent backcrossing might be used to transfer high seed weight into the high-yielding small-seeded lines. Singh (1977) suggested from correlation and path analysis studies on lentil that either phenotypic or genotypic correlations might be used in path analysis with equal efficiency. Positive association of harvest index with grain yield was observed. Grain yield and plant dry matter showed positive correlation with pod number, plant height, and number of primary and secondary branches but negative correlation with 100-seed weight. Dixit and Dubey (1984) observed strong direct effect of number of pods/plant on seed yield. Plant height and number of branches were the next highest contributors towards seed yield. Kumar and Sapra (1984) reported that yield/plant was directly linked with pod number/plant and pod size. Pods/plant was linked with plant height and the number of branches whereas time to flower and pod breadth negatively affected yield. Tyagi and Sharma (1985) observed that seed size had a highly significant negative correlation with seeds/pod. The traits of seeds/pod and total dry matter increased grain yield whereas harvest index negatively correlated with earliness. Balyan and Singh (1986) studying character association in lentil observed large effects of pods/plant and seed weight on seed yield. The relationship between seed yield and other component characters suggested that selection criteria based on component characters should be given due emphasis to exploit maximum yield potential in lentil. Mia et al. (1986) reported that seed yield/plant had positive correlation with plant height but negative association with time to flowering and seed size inferring that selection based on taller plant type with small seed size would be effective for high lentil seed yield. Manara and Manara (1988) observed positive correlation of seed yield was with number of pods/plant, number of one-seeded pods/plant, number of two-seeded pods/plant, number of empty pods/plant, number of days from seeding to maturity,
9
number of days from seeding to beginning of flowering, and number of days from flowering to maturity suggesting that seed yield could be increased by selecting later maturing plants with a greater number of pods. Baidya et al. (1988) suggested that short plant height with short growth period, without impairment of 100-seed weight might be considered as selection criterion lentil. Greater portion of total variance was observed due to environment. Seed weight/plot had the highest phenotypic, genotypic and environmental coefficient of variability. Heritability was highest for days to flowering, but its genetic advance was low. These studies indicated the advantages of upgrading lentil genotypes through simultaneous selection for plant height, dry weight/plant and number of pods/plant. Rao and Yadav (1988) observed that that selection based on harvest index, biological yield, and seed yield would improve lentil yield. Ramgiry et al. (1989) observed high heritability estimates for plant height, number of branches/plant, and harvest index whereas pods, seed weight and yield/plant showed low heritabilities. Correlation studies indicated that profusely branched plants with a high pod number had a high yield potential (Zaman et al. 1989). Jain et al. (1991) suggested that a combination of two or three variables, viz. plant height, branches/plant, and pods/plant was found better than other combinations of the characters for the improvement of seed yield in lentil. Hamdi et al. (1991) reported similar phenotypic correlations over two seasons, contrasting in rainfall, in the world lentil collection and similar genetic and phenotypic correlations were also observed over environments. Seed yield was positively correlated with straw yield, indicating that selection for either character will increase the other trait. Sharma et al. (1993) observed highly significant correlation between pod size and leaf size. Therefore, leaf size could be used as a fairly reliable index for the selection of seed size in segregating population, even at the seedling stage. The studies also revealed that number of seeds/pod had a strong negative correlation with 1000-seed weight, pod size,
10
and leaf area. Dutta et al. (1993) reported that reduced yields were due to shorter crop duration because of reduced sink sites, not to impaired grain development during high temperatures in the later part of the season. Esmail et al. (1994) reported negative correlation of time to maturity with seed yield/plant in two lentil crosses. Seed yield was positively correlated with branching, pods/plant, seeds/pod, seed weight and biological yield/plant. Branching, pods/plant and seeds/pod were observed to be the most important characters affecting yield variation. Kaiser (1994) studying stress tolerance to improve productivity for a target level of stress observed that in index selection theory the selection in non-stress environments would be more effective than direct selection for productivity under stress whenever the correlation between the two types of environments exceeded the heritability of productivity under stress. With high genetic correlation, selection should be conducted within a level of stress that maximizes heritability. In cases where heritability under nonstress is much higher than under stress, an index combining data from stress and non-stress environments is expected to be more efficient than selection based on evaluation only within stress environments. Secondary traits will be useful in breeding for productivity under stress whenever they have high heritability and high genetic correlation with productivity under stress. Jain et al. (1995) observed wide range of genetic variation in 21 lentil genotypes for plant height, number of branches/plant, biological yield, number of pods/plant, pod weight/plant, number of seeds/plant, seed index and harvest index. Heritability estimates were high for all traits except plant height and biological yield. There were significant positive correlation coefficients for all character combinations except seed index, which had negative correlation coefficients with the other traits. Path coefficient analysis revealed direct contribution of seed yield and seed index towards harvest index, while harvest index and biological yield contributed towards seed yield. Begum and Begum (1996) reported
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significantly positive correlations of seed yield with plant height, number of pods/plant, number of branches/plant and biological yield/plant. Positive relations of number of pods/plant with plant height, branches/plant and biological yield/plant were significant. Negative associations were found between seeds/pod and flowering time, seeds/pod and 100-seed weight, and branches/plant and 100-seed weight. It was suggested that plant height, number of branches and number of pods might be important criteria for lentil improvement. Bejiga et al. 1996 reported consistent regional differences among lentil landraces for time to flowering and maturity, 100-seed weight, seeds/pod and plant height. Erskine (1996) studied the seed-size effects on lentil yield potential and adaptation to temperature and rainfall in West Asia. Large-seeded material consistently had a longer reproductive growth period than the small-seeded group by 2.8 days, an extended period being required to fill its greater seed mass per pod. In the breeding material there was an advantage in seed yield of the large-seeded group over the small seeds in average temperatures < 10°C from January to April, with the converse true at higher temperatures. In the breeding material, the large-seeded group showed an advantage in seed yield over their small-seeded counterparts at the two wetter sites, whereas the small-seeded group was better adapted to dry environments. Correlation studies indicated the advantages of upgrading lentil genotypes through simultaneous selection for plant height, dry weight/plant and number of pods/plant. Abo-Shetaia et al. (1997) reported highly significant positive association between seed yield and seed index at the Giza environment whereas at Bostan environment, seed yield/plant was positively correlated with pods/plant, number of seeds/plant and negatively with number of seeds/pod. The use of these characters as selection criteria for seed yield was recommended. Dutta and Mondal (1998) reported that total dry matter, crop growth rate, net assimilation rate, relative growth rate, soluble protein content and harvest index showed positive
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associations with seed yield indicating prime importance of photosynthetic potentials of varieties and proper distribution of assimilates towards the reproductive sink for maximal yield expression. Kusmenoglu and Muehlbauer (1998) studying genetic variation for biomass and residue production in lentil noticed that plant height could be used for indirect selection for increased biomass. Seed yield might also be improved by selecting for larger vegetative biomass while keeping harvest index constant. Vir et al. (1998) reported a wide range of variability at both fertility levels in twenty-eight true breeding lines of lentil. High phenotypic and genotypic coefficients of variation (PCV and GCV) were observed at both fertility levels for seed yield/plant, biological yield/plant, pods/node, fertile nodes/plant, 100-seed weight and crushing hardness. High heritability at both fertility levels was estimated for biological yield/plant, harvest index, pods/plant, pods/node, fertile nodes/plant, days to flowering and maturity, and crushing hardness. The estimated expected genetic advance was high for seed yield/plant, biological yield/plant, harvest index, pods/plant, pods/node, 100-seed weight and crushing hardness at both fertility levels. Correlations, regression, path analysis estimates and per cent association of different traits with yield showed that in the irrigated crop, seed yield was mainly affected by pods and seeds/plant followed by total biological yield and harvest index. However, under rainfed conditions, seed yield largely correlated with 100-seed weight followed by harvest index, total biological yield and seeds/plant (Bhattacharya, 1999). Ferguson and Robertson (1999) observed that phenological adaptation, through sensitivity to photoperiod, temperature or both, appeared to be a major evolutionary force in wild lentils. Khattab (1999) observed that pods/plant, seeds/plant and 100-seed weight were the most important characters that contributed to grain yield. On the other hand, plant height, branching and pods/plant had the highest contribution on biological yield. Grain yield showed the highest positive direct effect on harvest index while biological
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yield had a negative direct and positive indirect effect via seed yield. Singh et al. (1999) reported that genotypic and phenotypic correlations exhibited similar trends but genotypic correlations were of higher magnitude than phenotypic ones. Days to 50 % flowering and clusters/plant showed higher estimates of heritability along with genetic advance. Days to 50 % flowering, clusters/plant, pods/cluster, seeds/pod and harvest index showed significant positive correlations with seed yield. Chakraborty and Haque (2000) observed that pods per plant and seeds per pod with a direct positive and negative effect of days to flowering and days to maturity, respectively, were the most important traits contributing to grain yield per plant. Tambal et al. (2000) in two lentil genotypes observed that although broad-sense heritability (h2) of the number of pods per inflorescence and its phenotypic correlation with seed yield were higher, the highest-yielding genotypes were not those with the most pods per inflorescence. Selection for the number of pods per inflorescence could not be recommended for increasing seed yield in lentil. Chauhan and Singh (2001) observed positive correlation of seed yield with number of secondary branches per plant, plant spread, number of fruiting nodes per plant and total biological yield per fruit. In general, these traits were also strongly correlated with each other. Safaei (2001) observed that 0.745 (R2) of the changes in yield were related to three characters (the number of pods, size of leaflet and 1000-seed weight). Path analysis showed that the direct effect of leaflet size on the yield was more than that of the other characters. Vir et al. (2001) studies comprising of 28 lentil genotypes showed positive and significant correlation of seed yield with harvest index, pods per plant, plant height, fertile nodes per plant and 100-seed weight. Path analysis revealed large direct effects harvest index, biological yield per plant, fertile nodes per plant and days to maturity on seed yield. Therefore, selection for harvest index, biological yield per plant, fertile nodes per plant and delayed maturity could enhance the seed yield. Studies of Kishore and Gupta (2002)
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revealed sufficient genetic variability for seed yield/plant, biological yield/plant, 100-seed weight, harvest index, seeds/pod, days to 50 % flowering, and days to maturity in crosses involving microsperma and macrosperma types. Biological yield/plant and seed yield/plant showed high degree of PCV, GCV and genetic advance. Heritability was high for biological yield/plant and days to maturity whereas moderate for seed yield/plant and most of the traits. Seed yield/plant showed significant positive association with biological yield/plant, harvest index, seeds/pod, 100-seed weight and days to 50 % flowering. Rathi et al. (2002) observed considerable closeness between genotypic and phenotypic coefficients of variation (GCV and PCV) indicating low influence of environmental factors in lentil. GCV and PCV were low for days to flower, days to maturity and protein content. Contrary to this, number of primary branches per plant, number of secondary branches per plant, number of clusters per plant, 1000-grain weight and grain yield per plant in both the generations (F1 and F2) and number of pods per cluster in F1 expressed high GCV and PCV but moderate heritab`ilities. Sinha and Singh (2002) reported that seed yield per plant was strongly and positively associated with number of seed and number of pods per plant. Seed yield was significantly correlated with plant height, and seed per pod. Naji et al. (2003) observed in exotic lentil cultivars that grain yield correlated positively with biological yield, plant height, yield per plant, seed per pod, pods per plant, and number of primary branches. Significant genotype x environment interaction was found for three growth rates and for other traits. Highly significant positive correlation was found between early growth rate per month and final biomass and seed yield, whereas nonsignificant and negative association was found between grain yield and days to flowering with the early line producing the highest yield. Sarker et al. (2003) reported that plant height, days to maturity and duration of flowering were the most important morphological and phenological traits contributing significantly to seed
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and straw yields in lentil. Singh et al. (2003) observed that positive and significant correlation of grain yield per plant with number of pods per plant and 100-grain weight. Number of fruiting branches, number of pods per plant and 100-grain weight had moderate to high positive direct effect on seed yield per plant. Bicer and Sakar (2004a and b) noted that the highest genetic variation was recorded for biological yield, grain yield and seed yield per plant. The highest heritability was recorded for seed weight and number of 50 % days to flowering. In another study, genotype x location interactions for biological yield per plant, seed yield per plant, number of pods per plant and number of seeds per plant were observed significant, and for these characters heritability was found low due to high environmental effects. Days to 50 % flowering, days to maturity and seed weight appeared to be useful traits because of high heritability. Kumar et al. (2004b) observed that harvest index, biological yield and days to 50 % flowering had the highest direct influence on lentil seed yield, suggesting the potential of these traits for utilization in selection programmes. Haddad (2004) noted significant variability for grain yield and yield related attributes between the sites where the lentil landraces were collected indicating specific adaptation of landraces to the local environment. Verma et al. (2004) observed that seed yield under rainfed conditions had highly significant positive correlations with number of pods per plant, number of seeds per plant, seed weight per plant, total dry matter/m2 and harvest index, whereas under irrigated condition, seed yield showed strong positive correlation with number of seeds per plant, total dry matter/m2 and harvest index. Although number of seeds per plant had significant correlation with seed yield, it had negative direct effect on seed yield under rainfed condition. Under irrigated condition, total dry matter/m2 had the highest positive direct effect on seed yield followed by harvest index. The indirect effects of total dry matter/m2 and harvest index through other characters on seed yield were either low or negligible. Sarker et al. (2005)
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studied seedling shoot and root characters in 8 lentil genotypes (collected from Ethiopia, India, Iran, Syria and ICARDA) during two seasons under field conditions. Combined analyses over two years showed that these characters exhibited significant genotypic variability. Stem length, taproot length and lateral root number were highly correlated, both amongst themselves and with yield. High heritability estimates provided reliability in screening based on these traits. Regression analysis showed that stem length alone accounted for 85 % of the variance that occurred in seed yield per plant. Cluster analysis showed that the landraces originated in Iran and Syria, and the breeding lines developed at ICARDA were distinctly different from the lentil accessions originated in countries at more southern latitudes (India and Ethiopia). Shrestha et al. (2005) observed that seed yield of the South Asian genotypes was higher than that obtained from the West Asian genotypes but lower than the crossbreds. The high seed yield of both the South Asian and crossbred genotypes was associated with rapid ground cover, early flowering and maturity, a long reproductive period, a greater number of seeds and pods, high total dry matter, greater harvest index, and high water use efficiency. West Asian genotypes, on the other hand, flowered later, matured later, and had a shorter reproductive period than the crossbred and South Asian genotypes. The greater seed yield of the crossbreds compared with the South Asian genotypes was the result of a similar increase in seed size (weight per seed). There were no significant differences in total root length, root dry matter, or water use among the 3 groups during the major part of the growing period. There was a significant difference in total water use due to the longer growing season of the West Asian genotype ILL 7983 and its ability to use late-season rainfall. Maximum water use efficiencies for seed yield of 7.0 kg ha-1 mm and for above-ground dry matter of 18.9 kg ha-1 mm were comparable with those reported in India and the Mediterranean environments of south-western Australia and Syria. Singh and Gupta (2005) observed
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positive and significant correlation of seed yield with number of primary and secondary branches per plant, pods per plant, plant height and biological yield showed under both early and late sown conditions in both years. Path analysis revealed that biological yield had the maximum direct effect on seed yield, followed by harvest index. All characters contributed indirectly to seed yield via biological yield. Thus, selection for high-yielding lentil genotypes should be based on primary and secondary branches per plant, pods per plant, biological yield and plant height. Studies of Andrews and McKenzie (2007) revealed that strategy to combat drought had been to match the crop’s development with the period of soil moisture availability. Lentil genotypes with early seedling establishment, early and rapid biomass development, and early flowering and maturity had been selected in sites of extremely low rainfall. In South Asia, lentil landraces exhibited a low diversity and discordance with landraces from other regions according to a combination of morphological characters (short or rudimentary tendrils and marked pubescence) and morpho-physiological characters (early maturity and low biological yield) (Erskine et al. 1989). This low diversity and discordance with other countries indicated a possible bottleneck (a temporary reduction in population size) when lentils were first introduced into South Asia (Erskine et al. 1998). El-Gawad et al. (1997) reported that seed and straw yields, plant height, number of branches, pods/plant, seeds/pod, seed index and crop and harvest index were greatly affected by edaphic and climatic conditions at two locations and three seasons. Jeena and Singh (2001b) studied 30 genotypes (28 wild accessions and two cultivars) of lentil using three separate hierarchical cluster analyses (viz., qualitative (HCA1), quantitative- (HCA2) and both qualitative and quantitative (HCA3) to work out the extent of genetic divergence on morphological and quality parameters. The results indicated wide genetic diversity as each analysis yielded formation of four, three and
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three clusters, respectively. The widest range of linkages i.e. Euclidian distances (Eds) were observed for cluster II followed by clusters I and III except in case of HCA2 where cluster I showed the highest range of Eds. Jeena and Singh (2002) reported that single seed weight, leaf rachis length and percentage germination were the characters contributing most towards genetic divergence among the lentil accessions. Kumar et al. (2004a) observed that genotypes of different eco-geographical regions clustered together. Number of days to 50 % flowering, number of clusters per plant, number of pods per cluster, number of pods per plant and seed yield per plant were found to be responsible for maximum differences between groups. Chauhan et al. (2005) found no parallelism between genetic and geographic diversity in lentil genotypes of heterogenous origin. Therefore, crosses between the members of clusters separated by high inter-cluster distances were likely to yield desirable segregates. The use of nuclear and biochemical markers (RFLPs, RAPDs, seed-protein electrophoresis) appeared to be the most consistent and reliable methods for determining genetic relationships. It is suggested that these techniques may be used in combination for taxonomic analysis of the genus Lens (Ahmad and McNeil, 1996). Abo-Elwafa et al. (1995) observed that genetic similarity among the genotypes was greater in population I (93.78 %) than in population II (89.20 %) indicating the efficacy of random amplified polymorphic DNA-polymerase chain reaction for the differentiation of lentil cultivars and lines. Kraic et al. (1995) reported that seed protein patterns obtained by SDS-PAGE were very similar in the varieties tested and not usable to distinguish between them. Similar results were obtained with RFLP analyses of 6 varieties when barley rDNA was used as a probe, but RAPD polymorphism was detected among these varieties, which fell into 2 groups. Ahmad et al. (1996) observed that the level of intraspecific genetic variation in cultivated lentils is narrower than that in some wild species and transmission of genetic
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material in Lens interspecific hybrids is genotypically specific, as identified by the RAPD markers. Alvarez et al. (1998) studied variability of random amplified polymorphic DNA (RAPD) markers in seven Spanish landraces and cultivars of Lens culinaris, including both macrosperma and microsperma types. Some polymorphism was observed within landraces, but not in cultivars. Three DNA bands distinguished macrosperma from microsperma accessions. Ford et al. (1997) using random amplified polymorphic DNA (RAPD) analysis observed polymorphism in all the lines with a maximum dissimilarity value of 0.36. This indicated a limited degree of genetic variation. Polymerase chain reaction (PCR) with primers based on the flanking regions of the 5S rRNA gene from Pisum sativum amplified the non-translated spacer (NTS) region from within the 5S rRNA gene of Lens. Three distinct amplification banding patterns differentiated between restricted genomic DNA of Lens spp. L. culinaris subsp. culinaris and L. culinaris subsp. orientalis shared similar markers of two distinctly different NTS sizes. L. nigricans and L. odemensis shared the same amplification pattern of a single sized NTS region. However, L. ervoides contained two separate sizes of NTS, distinct from other Lens species. In an effort to widen the genetic base of cultivated lentil, these species-specific molecular markers might be used to follow potential introgression between species. Rajora and Mahon (1997) examined mitochondrial DNA (mtDNA) and nuclear DNA (nuDNA) variation in 6 cultivars of L. culinaris subsp. culinaris and 2 accessions of L. culinaris subsp. orientalis. Each accession was clearly distinguishable from all others on the basis of both mtDNA and nuDNA fragment patterns. The mtDNA restriction fragment similarities ranged from 0.944 to 0.989, but nuDNA restriction fragment similarities varied from 0.582 to 0.987. Genetic relationships among accessions differed according to the source of DNA examined, although varieties Laird, Brewer and Redchief all exhibited high levels of mean similarity with both nuclear (0.982) and mitochondrial DNA (0.983).
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The studies of Ferguson et al. (1998a) revealed low diversity in landraces from South Asia (India, Nepal and Pakistan) and discordance with landraces from other countries using both isozyme electrophoresis and random amplified polymorphic DNA (RAPD) analyses. Germplasm from Afghanistan clustered with that from the South Asian countries. Based on genetic distance estimates from RAPD analysis, the countries with the lowest diversity were Pakistan, Afghanistan and Nepal. Zavodna et al. (2000) performed DNA polymorphism by analysis of variable number of tandem repeats (VNTR), (microsatellite and minisatellite DNA sequences), inter-simple sequence repeats (ISSR), and amplified fragment length polymorphism (AFLP) to distinguish between commercial lentil cultivars. DNA marker systems differentiated all the cultivars from each other. Rahman and Zafar (2001) studying the extent of genetic diversity among lentil cultivars through RAPD technique reported inter-varietal genetic relationships of the cultivars relating to their place of origin. Narrow genetic base in the local varieties was also proposed. Sonnante and Pignone (2001) observed differentiation of Ethiopian lentil landraces from the others in both cases (RAPD and ISSR analyses). On the other hand, Italian accessions showed a trend to group together. ISSR markers proved to be useful for distinguishing closely related genotypes, and possibly for substantiating the genetic peculiarity of ecotypes applying for the obtainment of origin and quality marks. Piergiovanni and Taranto (2003) observed that populations from Pakistan and Mediterranean countries were grouped in the same cluster, while those from Ethiopia showed the lowest homology with all tested countries based on seed storage protein profiles. Poonam (2006) observed that there was no parallelism between the two types of the clustering pattern using quantitative traits and RAPD primers. Yüzbaşıoğlu et al. (2006) using RAPD markers determined the genetic relationships among Turkish lentil cultivars and breeding lines. The dendrogram clearly showed two distinct groups. The
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first group was composed of Akm 563 and Akm 565. The second group contained all the cultivars and remaining breeding lines.
Genetic Studies on Quantitative Traits The subdivision of genetic variance into portions resulting from additive, dominance and epistasis effects of genes is very useful, since it provides information on the inheritance of quantitative characters and helps to identify appropriate breeding methods (Brim and Cockerham, 1961). Haddad et al. (1982) studied three lentil crosses and observed that additive genetic variance was the major component of variance in cross-2 (Tekoa x PI 212611) for all the characters except plant height and seed weight. Unexpectedly, estimates of dominance variance appeared to be high in crosses-1 (Chilean x PI 297784) and -3 (Precoz x PI 212611). Estimates of the additive x additive component seemed very small in all the three populations. The dominance component was consistently high for plant height in the three crosses and for seed weight in crosses-2 and -3. The unexpected high ratio of dominance variance to additive variance indicated the importance of this variance in these crosses. Bhajan et al. (1987) observed significant inbreeding depression in the F2 for yield/plant, test weight, number of pods, primary branches and secondary branches/plant. Crosses exhibiting high heterosis showed high inbreeding depression in the F2. El-Titi (1988) observed variable variation in the F2, F3, and F4 generations of lentil between crosses, and among generations with transgressive segregants in grain yield, 100-seed weight and secondary branches. The highest positive significant association was found between grain yield and number of seeds/plant followed by the association between grain yield and number of pods/plant. Tyagi and Sharma (1989) observed transgressive recombinants for extremely early flowering in crosses between the early flowering Precoz (ILL 4605), a moderately bold seeded macrosperma variety, and the early flowering Indian varieties L3991, JLS 3 and Sehore 74-3, in which
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early flowering was conferred by a gene(s) distinct from that (those) in Precoz. Waldia and Chhabra (1989) observed that both additive and non-additive components of variance were important for the number of branches/plant. They also observed that additive genetic variance was pronounced for pod number only. High heritability estimates were obtained for yield/plant, number of seeds/pod, secondary branches, branches/plant, and pods/plant. Singh and Singh (1990) studying lentil F1 to F3 generations observed that both GCA and SCA effects were significant in all generations, with a predominance of additive genes. Erskine et al. (1991) reported that overall trend for heterosis was noted to be greatest in hybrids with low yielding parents and least in those with high yielding parents. The problems of production of lentil hybrid seed, compounded with the generally low level of heterosis, rendered the development of hybrid lentil cultivars uneconomical for the foreseeable future. Sharma (1991) observed intermediate seed size in lentil F1 indicating incomplete dominance for seed size. Swarup et al. (1991) observed both D (variance due to additive effects of genes) and H1 (variance due to dominance effects of genes) components were significant in F1 generation of lentil crosses for time to flowering, plant height, and time to maturity suggesting the importance of both additive and dominance gene effects. The component H1 was significant and greater than D in magnitude for number of primary and secondary branches/plant, number of seeds/plant, number of pods/plant, number of seeds/pod, biological yield, seed yield, 100-seed weight, and harvest index indicating dominance gene action predominant in the expression of these traits. Studies of Chahota and Sharma (1993) observed significant differences for each character between lines and testers except for the number of primary branches, there were. Generally, the best specific combinations resulted from crosses between parents with high and low or both with low GCA effects. Singh and Singh (1993) reported the presence of both additive and non-additive components of genetic variance for 100-seed
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weight and seed yield. Kumar et al. (1994) observed that gene action was predominantly non-additive for protein and methionine content in lentil. Non-additive effects were relatively more important for seed yield and its components (Bajpai et al. 1995). The studies of Chauhan and Singh (1995a) showed that the magnitude of heterosis was affected by genotype x environment interaction. Heterosis for seed yield, early maturity, primary and secondary branches, pods/plant, seeds/pod and 100-seed weight was more distinct in cross combinations involving Precoz Sel. Chauhan and Singh (1995b) reported that multiple factors with incomplete dominance were responsible for seed size crosses between microsperma and macrosperma lentils. Chauhan and Singh (1995c) studying combining abilities, and both additive and non-additive gene actions observed that per se performance of parents gave fair indication of their GCA effects for 100-seed weight. Significant SCA effects for 100-seed weight were recorded in the crosses involving one parent with high and the other with low GCA effects. Chauhan and Singh (1995d) reported that additive and non-additive gene effects were important in the expression of protein content. Best specific crosses with high mean protein content may be exploited by harnessing additive and non-additive gene effects through bi-parental mating in early segregating generations for improvement of protein content. Tahir et al. (1995) observed additive gene action predominantly governed by recessive genes for the expression of seed weight suggesting simple selection in late generations when genes were fixed. Chauhan and Singh (1996) reported that both additive and non-additive genetic variances were important for all the traits studied suggesting pedigree selection and diallel selective mating systems of breeding for attaining maximum improvement in lentil. Gupta et al. (1996) reported that distant genetic inter-cluster distances represented the best opportunity for vigorous recombinants in lentil and deriving good combiners for better yield coupled with earliness. Predominantly additive gene action was reported for days to
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first flower, plant height, primary branches/plant and seeds/pod, and mainly non-additive for secondary branches/plant, pods/plant, 100-seed weight and yield/plant (Kumar et al. 1996). Chauhan and Singh (1997) noted preponderance of additive genetic variance for 100-seed weight and non-additive genetic variance for seed yield/plant in lentil. Yield/plant, secondary branches/plant, 100-seed weight, and biological yield/plant appeared to be useful traits because of high heritability and high genetic gain (Chauhan and Singh, 1998). Kant and Singh (1998) observed higher frequencies of transgressives for plant height in exotic x indigenous crosses whereas indigenous x indigenous crosses exhibited higher frequencies of transgressives for primary and secondary branches per plant. None of the crosses showed transgressive segregates for 100-seed weight in the F2 generation. Chuni et al. (2000) reported that genotypes of microsperma and macrosperma lentils, which possessed good general combining ability for complementary growth parameters, could be utilized in the hybridization programme to produce desirable genetic recombinants for further utilization. The crosses showing heterosis in both the F1 and F2 generations for leaf area and leaf area ratio, respectively, could be exploited through any simple selection method. The crosses revealing reduction of heterosis in F2 as compared to F1 were due to the epistasis interactions and were likely to produce transgressive segregants in the later generations. Chauhan and Singh (2001) observed environmental influence to the heterotic response for various traits in crosses between six microsperma types and one macrosperma type along with their F1s, including reciprocals. Rathi and Kumar (2001) observed positive heterosis for all the traits with maximum heterosis of yield, while the hybrids showing negative heterosis for either test weight or number of pods per cluster showed lower heterosis for yield. When heterosis was negative for both traits, a yield reduction was observed. Heterosis for yield had a positive association with the vigour of the component traits in lentil. Kahraman et al. (2004) determined the
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inheritance and heritability of winter hardiness in ten F6 derived recombinant inbred lines (RILs). Heritability estimates among the 10 RIL populations ranged from 15.9 to 90.7 %. Inheritance patterns of winter hardiness appeared to be quantitative on the basis of frequency distributions and the lack of discrete segregation classes. The dominance variance (δ2D) was observed high for harvest index, number of primary branches per plant, number of secondary branches per plant, and number of seeds per pod, while the same estimates attained a negative value for 100-seed weight. The additive genetic variance (δ2A) was also high for all the characters except for number of seeds per pod and number of primary branches per plant. The heritability in the narrow sense was high for all the characters except number of days to 50 % flowering and harvest index. Full-sib family selection would be suitable for all the characters studied except for number of days to 50 % flowering and harvest index (Singh and Singh, 2004). Studies regarding introgression of alien chromatin from wild Lens subspecies to cultivated lentil (Gupta and Sharma, 2007) revealed heterosis for plant height and branched/plant, and transgressive segregates for seed yield/plant also did appear indicating that wild subspecies can be exploited for breeding purposes and their variation can easily be utilized to widen the genetic base of the cultivated lentil.
Genetic Studies on Qualitative Traits During domestication and evolution, crop species underwent dramatic morphological and physiological changes. Genetic analysis of the control of these changes indicated that traits related to domestication were controlled by relatively few genes and that these genes often had major effects on plant phenotype. Moreover, in some species these domestication traits were clustered within the genome. Domestication might have occurred quite rapidly in some species as fixation of only a few genomic regions might have been sufficient for adaptation. Comparative mapping indicated that several
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genes for traits including plant height, flowering time, inflorescence and fruit morphology, organ color, and seed size and dispersal were conserved among different domesticated species. The limited number and apparently conserved nature of the genes controlling domestication facilitated the genetic manipulation of the related traits in both model and non-model crop species (Frary and Doĝanlar, 2003). To evaluate genetic variability and develop effective crop breeding systems, plant breeders need to have a definitive identification both of cultivars and selections of crop plants (Kjeldgaard and Marsh, 1994). Accurate identification of genotypes or varieties is very useful during all the steps of breeding from initial parent selection to the final utilization of cultivars in the production schemes (UPOV, 1991). Ladizinsky (1985) observed that in crosses between the cultivated species L. culinaris and its wild progenitor L. orientalis, a single recessive gene in homozygous condition controlled the hard seed coat of the wild species. In a cross between the wild species L. ervoides and L. culinaris, a single dominant gene controlled the hard seed coat of L. ervoides. Perretant (1994) reported complex inheritance of stipule shape while height and seed size both varied more in the F2 and F3 than in the parents. Ladizinsky (1997b) observed that two complementary dominant genes in intra- and inter-specific hybridization of wild and cultivated Lens species controlled dwarf phenotypes. Vandenberg and Slinkard (1990) reported that seed coat colour was determined by two independent loci. The dominant allele at one locus (Ggc) determined gray seed coat color, and the dominant allele at the second locus (Tgc) determined tan seed coat colour. The two dominant alleles (Ggc Tgc) interacted to produce a brown seed coat, whereas the double recessive (ggc tgc) was green. Seed coat pattern was determined by a series of five alleles at another locus. Marbled-1 (Scp(m1)) was dominant to marbled-2 (Scp(m2)), spotted (Scp(s)), dotted (Scp(d)), and absent (scp). Scp(m2) was dominant to Scp(s), Scp(d), and scp. Scp(s) and Scp(d) were codominant
27
alleles, both of which were dominant to scp. Malaviya and Shukla (1990) studying eight crosses between microsperma and macrosperma types, noted that flower and cotyledon colours were controlled by a single dominant gene. The seed coat colour showed a large number of intermediate colour combinations and was thought to be controlled by at least two pairs of genes. Singh and Singh (1992) reported that the F1 hybrids (crosses made between four grey mottled seed coat colour and one brown seed coat colour lentil lines) were brown seeded in all the crosses. Segregation pattern for seed coat colour revealed the control of a single dominant gene while a recessive gene was responsible for grey mottled seed coat colour. Inheritance studies of flower colour in lentil crosses of the violet x white crosses showed violet flowers in the F1 progeny, and the ratio of violet to white flowers in the F2 was 3:1, suggesting that violet flower colour was governed by a single dominant gene. This was supported by F3 segregation data (Singh and Singh, 1995). Emami and Sharma (1996b) confirmed digenic control of cotyledon colour in the F3 segregation of five crosses between parents with orange and light green cotyledons. Four crosses made between F2 plants of different crosses were raised from seeds with yellow and brown cotyledons. All F1 seeds had orange cotyledons. The F2 seeds harvested from F1 plants raised in a phytotron were analyzed immediately for cotyledon colour. The F2 seeds segregated in the ratio of 9 orange: 3 yellow: 3 brown: 1 green. The results from these crosses also confirmed digenic control of cotyledon colour in lentil. Emami and Sharma (1996c) studied inheritance of brown leaf pigmentation in eleven lentil crosses. Analysis of F2 plants revealed perfect monogenic segregation into 3 brown: 1 green ratio. Emami and Sharma (1996a) reported two cotyledon colours in lentil caused by dominant state of two unlinked genes, designated as Y (for yellow) and B (for brownish or dirty yellow). Double dominant condition YB gave orange (or red) cotyledons, while double recessive state by yybb produced light green cotyledons. Emami and Sharma (1999)
28
analyzed thirteen monogenic morphological marker traits of lentil for their joint inheritance in 25 crosses. Only three traits showed linkage: Ert (spreading-erect growth habit), Gs (brown-green stem) and Bl (brown-green leaf). The recombination frequency between Bl and Gs was 14 %, between Bl and Ert 34 %, and between Gs and Ert 38 %. Thus, the gene Bl is located between Ert and Gs. Khoddambashi and Sharma (1999) reported that the dark-green cotyledon colour was monogenic and the light green showed digenic inheritance. The involvement of three genes (Dg, Y and B) in the inheritance of cotyledon colour was considered. At the dominant state of gene Dg, the genes Y and B produced yellow and brown pigments, respectively. At the recessive state (dg dg), no pigment would be produced and the dark-green colour would appear. If gene Dg acted normally (dominant state), but both Y and B genes were at recessive state (Dg-yybb), again no pigment would be produced and cotyledons would be of light-green colour. Emami and Sharma (2000) studying the inheritance of testa (seed coat) colour and interaction of cotyledon and testa colours in lentil observed from the analysis of F2 and F3 seeds harvested from F1 and F2 plants respectively, that although black testa were dominant over non-black testa, its penetrance was not complete since both F1 plants and heterozygous F2 plants produced varying proportions of seeds with either black or nonblack testa. The F2 populations of the crosses between parents with brown and tan, as well as brown and green, testa segregated in the ratio of 3 brown: 1 tan and 3 brown: 1 green, respectively, indicating monogenic dominance of brown testa colour over tan or green. The expression of testa colour was influenced by cotyledon colour when parents with brown or green testa were crossed with those having orange or green cotyledons. Thus F2 seeds from these crosses with a green testa always had green cotyledons and never orange cotyledons. F2 seeds from these crosses with a brown testa always had orange cotyledons and never green cotyledons. Sharma and Emami (2002) reported a new gene which
29
functioned as a master gene for synthesis of the pigments determining cotyledon colour in lentil. This gene was different from the two earlier reported genes that were responsible for synthesis of yellow (gene Y) and brown (gene B) pigments. Double recessive homozygous condition of these two genes resulted into loss of both pigments and, consequently produced light green cotyledons. The new gene, in contrast, produced dark green cotyledons in recessive condition irrespective of the dominance or recessive state of the Y and B genes. It was hypothesized that the new gene for dark green cotyledon colour (Dg) acted at an earlier stage in the biosynthesis of the two cotyledon-specific pigments, which were derived from a common precursor, whose synthesis was blocked when Dg mutated to its recessive condition. Observations for leaf colour in lentil revealed a segregation ratio of 3 (dark green): 1 (light green). The F3 family data also fitted well to the ratio of 1 (dark green): 2 (segregating): 1 (light green) leaf colour. Individual F2 plants for plant pubescence segregated in the ratio of 3 (pubescent): 1 (non-pubescent). The F3 family data showed a ratio of 1 (pubescent): 2 (segregating): 1 (non-pubescent). The results revealed the involvement of single dominant gene for both the characters and gene symbols Dgl and Pub were proposed for them (Hoque et al. 2002). Kumar et al. (2005) studied monogenic inheritance and linkages for leaf colour, plant pubescence, number of leaflets per leaf, and plant height under field conditions. Normal green colour of foliage was found to be dominant over light green, pubescent plant over glabrous, high number of leaflets per leaf over low number of leaflets, and tall plant over dwarf. Linkage was estimated from joint segregation analysis, taking two characters at a time in all possible combinations. Gene symbols Gl, Pub, Ph, and Hl were proposed for these four traits, respectively. The genes were arranged in the order of Ph-Gl-Pub-Hl with the map distances of 21.1, 28.9, and 17.5 cM between them. Kant and Singh (1997) studied F2/F3 populations generated from crosses of exotic lentil with Indian cultivars under timely- and
30
late-sown conditions. Transgressive segregants for earliness were observed. Sarker et al. (1999a) reported single gene inheritance and polygenic system to control days to flower in lentil. Early flowering was determined by a single recessive gene (sn). Early flowering transgressive segregants occurred in F2 populations due to the interaction of sn and minor genes for earliness. Pubescent peduncle (Pep) was inherited as a single gene dominant to glabrous peduncle (pep). Tendrilless leaf (tnl) was controlled by a single gene recessive to tendrilled leaf (Tnl). The Sn, Scp (seed coat pattern), and Pep loci were linked together in linkage group 5 and Tnl was linked with Gs (coloured stem) in linkage group 1 of the lentil genome. Lentil landraces from South Asia exhibit low diversity and discordance with landraces from other countries and exhibited specific phenological adaptation which precluded the direct use of alien germplasm in breeding programmes. An understanding of the genetic relationships and diversity of South Asian lentil, in relation to lentil from other countries, is important in attempting to widen the genetic base of germplasm in the region (Ferguson et al. 1998a). Significant associations have been observed between quantitative characters and latitude of origin in lentil (Ferguson and Robertson, 1999), and variations between the sites where the lentil landraces were collected indicating specific adaptations to the local environment (Haddad, 2004). Interest in morphological variation in food legume species is increasing as plant breeders search for new variants to satisfy the adaptation requirements from new or changing environments or the needs of new end-users. Traits affecting the development of the crop canopy or the seed, including for example photosynthate repartitions, could have an impact on yield, quality and diseases. Increased attention to these complex interactions through international multidisciplinary cooperation has contributed, and could further contribute, to progress in breeding and disease management, thus ultimately resulting in an improvement of the
31
yield potential, yield stability and quality of the crops (Porta-Puglia et al. 2000). Significant changes in morphological and phenological traits have been observed in regional adaptation, and the response to different selection methods largely depends on landrace and selection site (Horneburg and Becker, 2008). The identification and/or development of cultivars which are phenologically well-adapted to their environment is, therefore, likely to contribute substantially to future yield improvements (Whitehead et al. 1998). The search to develop a productive "plant type" for major agro-ecological zone to serve as a platform for local adaptation, and then subsequently breed for location-relevant traits such as abiotic stresses, resistance to diseases and pests may be imperative (Evenson and Gollin, 2003) for lentil improvements. In countries such as Pakistan and India, lentil availability is little more than 1 to 2 g day−1 (FAOSTAT, 2007) although Indian subcontinent is the largest lentil producing region in the world, contributing about 42% of the total world production (Mishra et al. 2007). In Pakistan, lentil is planted on an area of 33.9 thousand hectares annually producing 17.9 thousand tonnes with an average yield of 528 kg ha–1 (Agricultural Statistics of Pakistan, 2006). The average yield is low and the key to increasing yield potential is through widening the available genetic base. Seed yield is the primary objective in most of the lentil breeding programmes. However, under certain situations other traits may have an equal or greater importance. Armed with an understanding of the specific adaptation of lentil, the local constraints to production and various consumers’ requirements of the region for seed, the information regarding associations among yield and yield related traits with special reference to the phenology, diversity in the germplasm to exploit the genetic variation and identify desirable germplasm for use as parentals, detection and estimating the components of genetic variation (additive, dominance and epistasis) for yield and yield contributing traits in the segregating populations, and
32
determining the pattern of gene action for important qualitative traits to select desirable recombinants with stable and improved seed yield are the essential aspects. Thus, knowledge of the genetic control of characters is of considerable importance in organizing plant breeding programme for the improvement of the concerned crop plant. The present studies, therefore, were planned to assess the genetic variation and determine the relationships among different plant/yield related traits in the germplasm, genetic diversity in the parental lines used for the production of segregating populations for the nature of gene actions in quantitative and pattern of inheritance in qualitative traits. The information gained may contribute to elaborate selection strategies in lentil breeding and help increase the productivity of lentil in the development of varieties possessing improved plant phenology and yield related traits.
33
Chapter 2
MATERIALS AND METHODS In the present studies, 117 lentil germplasm genotypes received from the International Center for Agricultural Research in the Dry Areas (ICARDA), Syria belonging to different countries throughout the world including Pakistan were evaluated to assess the genetic variation under the prevailing local climatic/edaphic conditions and to find out the possibilities of seed yield improvements in lentil through the estimation of relationships among different plant and yield related traits, and partitioning their direct and indirect effects as well as by calculating different genetic parameters. Estimates of genetic diversity in the parental lines used for the production of segregating populations in the inheritance studies were also determined through morphological characters as well as by molecular markers. For molecular studies, randomly amplified polymorphism DNA (RAPD) technique was used. Parents were selected on the basis of their morphological dissimilarities they possessed for their phenology, growth habit and seed characteristics. The populations derived from hybridization were subjected to biometrical analysis for estimation of nature of inheritance for different plant/yield related traits. These populations consisted of parents, their F1s, backcrosses with either of the parent (BC1s and BC2s) and the F2s. The inheritance studies on some of the qualitative plant traits were also carried out to assess the pattern of inheritance in these traits for their use as markers in the future breeding programme. Materials and methods involved in each experiment varied, hence each experiment is described accordingly.
34
2.1 Assessment of Genetic Variation in Lentil Germplasm The germplasm was comprised of 117 diverse genotypes received from the International Center for Agricultural Research in the Dry Areas (ICARDA), Syria including locally developed/selected lentil varieties. The experiment was conducted at the Nuclear Institute for Agriculture and Biology (NIAB), Faisalabad (31.3 oN and 73.10 oE), Pakistan following randomized complete block design (RCBD) with three replications maintaining plant to row distance of 10 x 30 cm with 4 meter long row. Soil moisture was preserved by irrigating the land with canal water prior to sowings. The experiment was sown in extensive moisture/wet conditions during 1st week of November, 2002. Recommended dose of fertilizer (DAP fertilizer @ 120 kg ha-1) was applied at the time of sowing. Out of 117, twenty-five genotypes did not flower under Faisalabad conditions and data on 92 genotypes were recorded for different characters. The details of the germplasm comprising of 92 genotypes and meteorological data for the growing season of the crop are given in tables 2.1 and 2.2 respectively. The data for days to flowering (50 %) were recorded on line basis. The calculations were made from the day of sowing. Other traits were recorded when plants were physiologically matured. Data on five randomly selected plants were recorded for plant height (cm), primary branches per plant, secondary branches per plant, nodes and fertile nodes on primary branch, nodes and fertile nodes on secondary branch, seeds per pod (mean of five pods from each plant), pods per plant, pod length and pod breadth in centimeters (mean of five pods from each plant), 100-seed weight (g), biomass per plant (g), harvest index per plant and seed yield per plant (g). Analysis of variance was employed to test the significance among genotypes following Steel and Torrie (1984) using computer software “MStat C” for Windows.
35
Table 2.1: Details of lentil germplasm evaluated at the Nuclear Institute for Agriculture and Biology (NIAB), Faisalabad, Pakistan during 2002-03 S. No. 01 02 03 04 05 06 07 08 09 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46
Genotype ILL 292 ILL 268 ILL 4679 ILL 4605 ILL 515 ILL 4411 ILL 7789 ILL 5888 ILL 4793 ILL 8591 ILL 6968 ILL 512 ILL 4737 ILL 75 ILL 8385 ILL 69 ILL 8594 ILL 236 ILL 788 ILL 1920 ILL 1672 ILL 4740 ILL 103 ILL 8368 ILL 1921 ILL 494 ILL 2465 ILL 2580 ILL 7556 ILL 7715 ILL 8117 Pant L 406 Pant L 639 ILL 604 ILL 750 ILL 910 ILL 55 ILL 341 ILL 68 ILL 637 ILL 2275 ILL 4804 ILL 8411 ILL 8412 ILL 622 ILL 495
Source Algeria Argentina Argentina Argentina Azerbaijan Bangladesh Bangladesh Bangladesh Belarus Bolivia Brazil Bulgaria Canada Chile China Cyprus Ecuador Egypt Egypt Egypt Ethiopia France Germany Georgia Great Britain Guatemala India India India India India India India Iran Iran Iran Iraq Italy Lebanon Lebanon Lebanon Libya Libya Libya Macedonia Mexico
S. No. 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92
36
Genotype ILL 97 ILL 4609 ILL 4782 ILL 4606 ILL 2189 ILL 4404 Masoor 93 NL 2002 Turk Masoor ILL 8499 ILL 4803 ILL 6845 ILL 4922 ILL 84 ILL 7744 ILL 79 ILL 1861 ILL 5505 ILL 26 ILL 1867 ILL 2130 ILL 4400 ILL 4401 ILL 4525 ILL 5748 ILL 5782 ILL 6024 ILL 6463 ILL 6468 ILL 6821 ILL 6126 ILL 507 ILL 1890 ILL 107 ILL 590 ILL 962 ILL 1513 ILL 8257 ILL 82 ILL 4778 ILL 4875 ILL 225 ILL 950 ILL 4741 ILL 4763 ILL 1933
Source Morocco Netherlands Norway Palestine Pakistan Pakistan Pakistan Pakistan Pakistan Peru Poland Poland Portugal Russia Saudi Arabia Spain Sudan Sudan Syria Syria Syria Syria Syria Syria Syria Syria Syria Syria Syria Syria Tajikistan Tunisia Tunisia Turkey Turkey Turkey Turkey Turkmenistan Ukraine Uruguay Uzbekistan Yemen Yemen Yemen Yemen Yugoslavia
Table 2.2: Meteorological data for the growing season of lentil germplasm during 2002-03 Month
Air Temperature (oC) Maximum Minimum
Relative Humidity (%)
November
Total Rain Fall High Low Mean High Low Mean High Mean Low Mean (mm) 30.8 22.8 27.7 17.6 6.5 12.2 93 80.4 27 45.3 T
December
26.8
17.0
22.8
16.0
3.1
8.1
94
84.4
35
49.7
2.0
January
22.8
8.8
16.5
11.0
0.6
4.8
100
92.8
42
69.7
7.6
February
26.6
17.3
22.0
15.5
3.5
8.7
100
86.5
33
53.6
76.2
March
32.6
17.0
26.7
20.2
4.5
13.5
93
74.8
32
49.2
38.4
April
41.4
28.8
35.7
24.5
10.8
19.9
78
54.2
20
36.5
-
T: Traces Source: Plant Physiology Section, Ayub Agricultural Research Institute, Faislalabad
37
Correlation and regression analyses were carried out, and partitioned into direct and indirect effects (Wright, 1921; Dewey and Lu, 1959). Principal component analysis (PCA) was performed to find out the traits accounting for as much of the phenotypic variation of the genotypes using computer software “Minitab 11” for Windows. For PCA, procedures of Chatfield and Collin (1980), Mahloch (1974), Mazlum (1994), and Mazlum et al. (1999) were followed. The component loadings (correlation coefficients) and the variances (eigenvalues) regarding the components were computed for all the characters at the first step following correlation matrix as all the characters had equal importance with different scales. The proportion of the total variance explained by each principal component was additive, with each new component contributing less than the preceding one to the explained variance. Subsequently, varimax rotations were made for these components to eliminate medium-range loadings (correlations) to make the interpretation of the components easier. Sixteen rotated components which were too numerous to explain and components which explained a relatively small proportion of the total variance of the principal components were eliminated for simplification. Then the first six components selected whose eigenvalue (λ) values were > 1, and were rotated. In the last step, the number of components considered was eight. Subsequently, these eight components were rotated. Since all the communalities were larger than 0.7 in this final case, it might be assumed that all the variables were described to an acceptable level and these eight components accounted for 86 % of the total variance of their original data. This showed that first eight components could be considered significant in the analysis. In general, component loadings (correlation coefficients) larger than 0.6 were taken into consideration. The components with larger variances were more desirable since they gave more information about the data.
38
2.2
Estimation of Genetic Diversity in Parental Lines To produce materials for investigation of nature of gene action applying
generation means analysis, twelve genotypes (ILL 4605, ILL 5782, ILL 6024, ILL 6821 and Turk Masoor-macrosperma types; and ILL 2580, ILL 6468, ILL 7556, ILL 7715, ILL 8117, Masoor 93 and Pant L 406-microsperma types) of diverse origin possessing contrasting phenological, growth, seed and other plant characteristics were selected. The genotypes ILL 2580, ILL 7556, ILL 7715, ILL 8117, Pant L 406 and Masoor 93 were well adapted to the growing conditions while the genotypes ILL 4605, ILL 6024, ILL 6468, and ILL 6821 (exotic material received from ICARDA) showed performance problems in this environment. The landrace Turk Masoor was collected from the market (purified for uniformity a few years ago). A detailed description of the parental lines used and meteorological data of the growing season are given in appendix I and II respectively.
2.2.1
Estimation of genetic diversity in parental lines using morphological characteristics
The seeds of 12 genotypes were sown in randomized complete block design (RCBD) with three replications keeping plant to row distance of 10 x 30 cm with 3 rows of 4 m long during 1st week of November 2001 at NIAB, Faisalabad. Meteorological data for the growing period of the crop is presented in appendix-III. Prior to sowings, soil moisture was preserved by irrigating the land with canal water. Recommended dose of DAP fertilizer @ 120 kg ha-1 was applied at the time of sowing. Other necessary cultural practices were carried out through out the growing season. The data for days to flowering (50 %) were recorded on line basis. Other traits were recorded when plants were physiologically matured. Data on five randomly selected plants were recorded for plant 39
height (cm), primary branches per plant, secondary branches per plant, fertile nodes on primary and secondary branches, pods per plant, 100-seed weight (g), biomass per plant (g), harvest index per plant and seed yield per plant (g). Data recorded were subjected to analysis of variance to test the significance among genotypes following Steel and Torrie (1984) using computer software “MStat C” for Windows. Principal component analysis (PCA) was performed to find out the traits accounting for as much of the phenotypic variation of the genotypes using computer software “Minitab 11” for Windows as described in section 2.1.
2.2.2
Estimation of genetic diversity in parental lines using randomly amplified polymorphic DNA (RAPD) markers
Isolation of plant genomic DNA The seeds of twelve genotypes were sown in Petri plates in the incubator at 22±3 o
C for 21 days. Young leaf samples (approx. 1g) were collected from the seedlings. The
samples were kept in polypropylene tubes with water until before grinding. Each sample was ground to a very fine powder in liquid nitrogen and transferred to a chilled (−20 ºC) 2 ml centrifuge tube. Then added 665 µl of extraction buffer without SDS and subjected the samples to Vortex to suspend avoid samples. Added 115µl of 5M NaCl, mixed well by gentle inversion until samples were evenly suspended, added 90 µl of 10 % CTAB (NCetyl, N, N, N-trimethylammoniumbromide) in 0.7M NaCl (10 % CTAB in 0.7M NaCl solution was prepared by taking 10g of CTAB plus 50 ml of 1M Tris HCl having pH 8.0 and 20 ml of 0.5M EDTA of pH 8.0. For 0.5M EDTA, add 8 g of NaOH in 93.06 g of EDTA in 500 ml H2O, adjust pH at 8.0, then sterlize by autoclaving). These were meshed up to 1000 ml with distilled water and then sterlized by autoclaving) and incubated for 10
40
minute at 65 0C. Added 900 µl Chloroform and mixed well, centrifuged at 12000 rpm (rotation per minute) for 2 minute. Collected the supernatant (upper layer), transferred in a new tube, added 600 µl isopropanol, spun down the fish out the DNA when visible strands were formed. Rinsed with 70 % ethanol and spun down at 12000 rpm for 2 minute and the DNA was air-dried for 5-10 minute. Added 50 µl of 1 X TE buffer (take 10mM Tris base of pH 8.0 and 1mM EDTA of 8.0 pH to adjust the volume to 100 ml dH2O, and sterlize by autoclaving), spun down at 12000 rpm for 1 minute, added 7 µl RNAs, mixed gently and incubate for 1 hour at 37 oC and kept in ice. 1µl sample was taken and 9 µl 1 X TE buffer was added to make 10 µl volume of the sample. After this treatment, in 1µl of each sample, 4 µl of blue Juice ((for 10 X loading buffer, take 65 % w/v sucrose, 10 mM Tris-HCl of pH 7.5, 10 mM EDTA and 0.3 % w/v Bromophenol Blue) were added for measuring the concentration of DNA. The quantity of the total DNA isolated from plants was measured by DyNA Quant 200 (Pharmacia Biotech) Fluorimeter using Hoechst 33258 dye and qualitatively by agarose gel electrophoresis using UV-visible spectrophotometer. The concentration used for RAPD analysis was adjusted to 10ng/μl. DNA samples were diluted in sterile distilled water to a concentration of 12.5 ng/µl for PCR analysis.
PCR reaction for RAPD Random decamer primers (Operon series) were dissolved in sterilized distilled water at a concentration of 25 ng/µl. Fifteen primers belonging to Operon kits (OPA-01, OPA-13, OPB-01, OPB-05, OPB-09, OPB-19, OPN-02, OPN-04, OPN-05, OPN-06, OPN-13, OPN-14, OPN-15, OPN-16, OPN-18) were employed (Appendix-II) for PCR amplification. Amplifications were carried out in a 25-µl reaction volume containing 2.50 µl of 10X PCR buffer {(100 mM Tris-HCl, pH 8.3 at 25 oC), (500 mM KCl, 0.01 % gelatin)}, 3.00 µl of 25 mM MgCl2, 2.00 µl of 10 mM dNTPs (10 mM each of dATP,
41
dGTP, dCTP, and dTTP), 0.25 µl of unit Taq DNA polymerase, 2.00 µl of 25 mM primer, 5.00 µl of 12.5 ng/µl-template DNA, 2.50 µl gelatin and 10.25 µl of sdH2O. The reaction mixture was overlaid with one drop of mineral oil to avoid evaporation. The amplifications were carried out in A Perkin Elmer Thermal Cycler (Eppendorf Master Cycler Gradient PCR system), programmed for denaturation steps of 5 minute at 94 oC and of 1 minute at 94 oC, annealing for 1 minute at 36 oC and 2 minute at 72 oC for primer extension, and then followed by 35 cycles up to 2nd step of denaturation, extension for 10 minute at 72 oC. After completion of all the steps, final extension step was kept at 72 oC for 5 minute and then held the PCR products at 4 oC until these (12 µl each sample) were separated on 1% agarose gel (add 3.5 g of agarose in 350 ml of 1 X TAE buffer. To get 350 ml of 1 X TAE buffer, add 7 ml of 50 X TAE plus 343 ml of sterile dH2O), stained with 5 µl of ethidium bromide, along with 1 kb ladder {(working solution (0.05 µg/µl) is dikuted from the stock (0.9 µg/µl) by mixing 11 µl of 1 kb DNA ladder stock with 20 µl of 10 X loading buffer and 169 µl of sterile dH2O and stor at 4 oC)} at 80-90 volts for 1 hour. After appropriate running of the dye, the power supply was disconnected and the electrophoresis gel was observed under UV transilluminator and RAPD amplification products were documented by Eagle Eye Gel Documentation system (Stratagente, USA).
RAPD analysis The PCR products were measured as polymorphic bands pattern and scored as either present or absent (bi-state). The data was used to estimate similarity on the basis of Jaccard’s method. Genetic relationships among the genotypes were measured following un-weighted pair group method of average (UPGMA) clustering algorithm (Nei and Li, 1979) which provides results most consistent with known heterotic groups and pedigree information (Kantety et al. 1995).
42
2.3
Estimation of Nature and Pattern of Inheritance of Quantitative and Qualitative Traits 2.3.1
Hybridization for production of F 1 , F2 , BC 1 and BC2 populations
Seeds of twelve selected genotypes were divided into two sets and each was sown during November 1999 at 15 days interval, at the Nuclear Institute for Agriculture and Biology (NIAB), Faisalabad, Pakistan. Each entry was sown in three rows keeping each row 4 meter long and maintaining plant to row distance of 10 x 30 cm apart. Emasculations were done manually in the morning with the help of forceps. The emasculated flowers were tagged. Next day each emasculated flower was pollinated. Each flower was pollinated two times to ensure the pollination success. The crosses were attempted in the following manner. S. No. Cross Combination 1. 2. 3. 4. 5. 6.
S. No.
Female x Male ILL 5782 x ILL 2580 ILL 2580 x ILL 5782 ILL 6024 x ILL 8117 ILL 8117 x ILL 6024 ILL 6468 x ILL 7556 ILL 7556 x ILL 6468
7. 8. 9. 10. 11. 12.
Cross Combination Female x Male ILL 6821 x ILL 7715 ILL 7715 x ILL 6821 Masoor 93 x Turk Masoor Turk Masoor x Masoor 93 ILL 4605 x Pant L 406 Pant L 406 x ILL 4605
At maturity, sufficient crossed materials along with their parents were harvested and stored for next sowings. As contrasting seed coat pattern and cotyledon colours were used as markers in the parental lines, the crossed seeds were easy to recognize for collection at the time of harvest. For producing BC1, BC2 and F2 generations, the harvested F1 seeds along with their respective parents were sown during November 2000 at the Nuclear Institute for 43
Agriculture and Biology (NIAB), Faisalabad. Back crosses (BC1: F1 x Female parent and BC2: F1 x Male parent) and fresh crosses were attempted using the same technique applied for producing F1 seeds. At maturity, the crossed seeds along with their respective parents were harvested for next sowing.
2.3.2
Raising populations for inheritance studies
In this experiment, the material consisted of parents (ILL 2580, ILL 4605, ILL 5782, ILL 6024, ILL 6468, ILL 6821, ILL 7556, ILL 7715, ILL 8117, Masoor 93, Pant L 406, and Turk Masoor), their F 1 s, F 2 s, BC 1 s and BC 2 s. The experiment was planted during 1 st week of November 2001 at the Nuclear Institute for Agriculture and Biology (NIAB), Faisalabad. Meteorological data for the growing period of the crop is presented in appendix-III. The soil type was sandy loam. Soil moisture was preserved by irrigating the land with canal water. At the time of field preparation, recommended dose of fertilizer (DAP @ 120 kg ha-1) before sowing was applied. The experiment was laid out following randomized complete block design (RCBD) with three replications maintaining plant to row distance of 10 x 30 cm with 4 meter long row. Seeds of F1, BC1 and BC2 generations of each cross were sown in a single row-plot. For F2 populations, ten rows of each cross and three rows of each parent in a replication were sown. One border row on either side of each block was treated as non-experimental and was sown with parental seeds. Other normal cultural practices were carried out throughout the growing season to maintain the experiment.
44
2.3.3
Data recording for estimation of nature of inheritance on quantitative traits and pattern of inheritance on qualitative traits, and statistical analyses
The data on different plant traits were recorded at appropriate stage from individual plot on plant basis during the whole crop period. The traits included days to flowering (from sowing to first flower), days to 90 % pod maturity, plant height (cm) at maturity, primary branches per plant, secondary branches per plant, nodes on primary branch, fertile nodes on primary branch, nodes on secondary branch, fertile nodes of secondary branch, seeds per pod, pods per plant, pod length (cm), pod breadth (cm), 100seed weight (g), biomass per plant (g), harvest index per plant and seed yield per plant (g). The data were recorded on twenty randomly selected plants in the P1, P2, BC1, and BC2 generations whereas data in F2 generations were recoded on one hundred plants in each replication. The data collected on the characters under study to estimate the nature of inheritance were subjected to statistical analysis following analysis of variance (Steel and Torrie, 1984). Significance of differences among all the generation means was tested according to Little and Hill (1978) as follows. Coefficients for partitioning of sum of squares among six generations into orthogonal comparisons Comparison* P1’s P2’s F1’s F2’s BC1’s BC2’s P1s vs P2s
1
–1
0
0
0
0
P’s vs F1s
1
1
–2
0
0
0
BC1s vs BC2s
0
0
0
0
1
–1
BC’s vs F2’s
0
0
0
2
–1
–1
P’s, F1’s vs BC’s, F2’s
1
1
1
–1
–1
–1
* P1s: Female parents, P2s: Male parents, P’s: Both female and male parents, F1s: 1st filial generations, BC1’s: Backcrosses with female parent, BC2’s: Backcrosses with male parent, F2s: 2nd filial generations
45
The formula is as under: SS = (Σci Yi /r Σci 2) Where; SS = Sum of squares of comparison Σ = Summation ci = Comparison coefficient Yi = Generation totals r = Replications To estimate the nature of inheritance in different quantitative plant/yield and yield related traits, the data was subjected to generation means analysis as described by Mather and Jinks (1982). Adequacy of scale must satisfy two conditions namely, additive effects of gene and independence of heritable components from non-heritable ones. The test of first condition provides information regarding absence of gene interactions. The test of adequacy of scales is important because components of variance are made assuming the absence of gene interaction. Joint scaling test (Cavalli, 1952) was performed to detect epistasis for each character. In the presence of epistasis, additive (d), dominance (h) effects, and non-allelic interactions (additive x additive (i), additive x dominance (j) and dominance x dominance (l) were estimated following Hayman (1958). For pattern of inheritance in qualitative traits such as seedling growth habit, epicotyls pigmentation, seed coat pattern and cotyledon colours, the data were recorded in parents, F1 and F2 generations. The seeds for cotyledon colours were analyzed by scratching some portion of the seed avoiding damage to embryo. The data collected were analyzed applying χ2 test.
46
Chapter 3
Genetic Variation in Lentil Germplasm Identification of genotypes for certain desirable traits has become increasingly important for genetic research and breeding. Genetic improvement in any quantitative trait depends upon effective selection among individuals that differ in phenotypic value. Effective selection is possible only when genetic variability existed among the populations to be explored. To utilize germplasm efficiently and effectively, it is important to investigate the genetic variation it contains. Many tools are now available for identifying desirable variation in the germplasm including total seed protein, isozymes and various types of DNA markers. However, morphological characterization is the first step in the description and classification of crop germplasm (Smith and Smith, 1989; Singh and Tripathi, 1989). The mean square, range and mean values for each character in 92 genotypes of lentil germplasm are presented in table 3.1. Highly significant differences were observed among the genotypes for all the characters under study. Coefficient of variation (C. V.) among the genotypes indicated that days to flower (1.53 %), seeds per pod (1.91 %), pod length (2.34 %) and breadth (2.96 %), 100 seed weight (3.65 %), harvest index (5.20 %), plant height (6.89 %), nodes on primary branch (7.49 %), pods per plant (7.62 %), biomass (8.51 %), fertile nodes on primary branch (8.63 %), primary branches (8.67 %) and secondary branches (8.80 %) had low to moderate variability. Greater variations were observed for fertile nodes on secondary branch (12.64 %), seed yield per plant (9.71
%)
and
nodes
on
secondary
branch
47
(9.11
%)
in
the
germplasm.
Table 3.1: Values of mean square, range, means and coefficient of variance for different plant characteristics in lentil germplasm Parameter
Mean Square 800.71**
69.37
134.70
116.87
C. V. (%) 1.53
112.33 **
28.08
48.77
37.24
6.89
0.10 **
1.82
3.12
2.63
8.67
72.56 **
8.91
33.13
18.95
8.80
48.14 **
13.67
32.85
23.41
7.49
42.92 **
4.80
20.30
12.71
8.63
36.11 **
12.57
26.74
19.01
9.11
35.55 **
4.04
20.50
9.45
12.64
0.09 **
1.00
1.67
1.18
2.34
Pod breadth (cm)
0.12 **
0.47
0.87
0.62
2.96
Seeds pod-1
0.15 **
1.04
1.93
1.62
1.91
Pods plant-1
13915.90 **
57.79
427.91
168.25
7.28
100 seed weight (g)
2.11 **
0.87
4.57
2.17
3.65
Biomass plant-1 (g)
81.73 **
3.81
26.52
14.69
8.51
149.28 **
12.02
39.39
25.10
5.20
10.17 **
1.01
8.15
3.82
9.71
Days to flower (50%) Plant height (cm) Primary branches plant-1 Secondary branches plant-1 Nodes on primary branch Fertile nodes on primary branch Nodes on secondary branch Fertile nodes on secondary branch Pod length (cm)
Harvest index plant-1 (%) Seed yield plant-1 (g)
C. V.: Coefficient of variance
Range
Mean
**: Significant at P < 0.01
48
Flowering among the genotypes ranged from 69.4 (NL 2002) to 134.7 days (ILL 8385). Plant height ranged from 28.1 (ILL 950) to 48.8 cm (ILL 79). Maximum primary branches (3.1) were noted in genotype ILL 6024 while ILL 6821 produced only 1.8. Genotype ILL 4922 produced maximum secondary branches (33.1) while minimum (8.9) were noted in ILL 1867. Nodes on primary branch ranged from 13.7 (ILL 4401) to 32.9 (ILL 4649). Genotype ILL 4778 produced maximum fertile nodes (20.3) on primary branch whereas in ILL 8385, only 4.8 fertile nodes were observed on primary branch. Genotype ILL 6845 produced maximum nodes on secondary branch (26.7) while minimum (12.6) were noted in ILL 622. Maximum number of fertile nodes on secondary branch (20.5) was noted in ILL 5505 whilst minimum number (4.0) was recorded in ILL 8412. The longest pod was observed in ILL 4400 (1.67 cm) while ILL 82 had short pods (1.00 cm). The pod breadth ranged from 0.50 cm (ILL 7744) to 0.87 cm (ILL 4679). Seeds per pod ranged from 1.04 (ILL 950) to 1.93 (ILL 1921). The genotype ILL 4404 produced maximum number of pods (427.9) whereas minimum (57.8) were recorded in ILL 4793. The largest seed size was observed in ILL 5748 (4.6 g per 100 seed weight) while ILL 84 had minimum seed weight of 0.9 g. The genotype ILL 2580 produced the highest biomass per plant (26.5 g) as compared with the lowest biomass of 3.8 g (ILL 4778). The highest harvest index (39.4 %) was recorded in ILL 4679 whereas ILL 1890 had the lowest harvest index (12.0 %). Seed yield per plant among the genotypes ranged from 1.0 g (ILL 4778) to 8.2 g (ILL 68). Correlation Coefficients: The results regarding genotypic and phenotypic correlation coefficients are given in table 3.2. The magnitudes of genotypic correlation coefficients for all the traits with seed yield per plant were slightly higher as that of their phenotypic counterparts. Seed yield per plant positively correlated with biomass per plant (rg = 0.708*, rp = 0.703**), pods per plant (rg = 0.483*, rp = 0.479**), 100 seed weight (rg = 0.313*, rp = 0.310**), fertile nodes on primary branch (rg = 0.249*, rp = 0.243*), harvest
49
Table 3.2: Genotypic (G) and phenotypic (P) correlations (r) among various plant characteristics in lentil germplasm Parameter
r
Plant Height
Days to flower
G P
-0.17546 -0.17053
Plant height
G P
Primary branches plant-1
G p
Secondary branches plant-1
G P
Nodes on primary branch
G P
Fertile nodes on primary branch
G P
Nodes on secondary branch
G P
Fertile nodes on secondary branch
G P
Pod length
G P
Pod breadth
G P
Seeds pod-1
G P
Pods plant-1
G P
100 seed weight
G P
Biomass plant-1
G P
Harvest index plant-1
G P
Primary branches plant-1 0.04251 0.03262
Secondary branches plant-1 -0.02514 -0.02592
Nodes on primary branch 0.05294 0.05109
Fertile nodes on primary branch -0.30757* -0.30263**
Nodes on secondary branch 0.03043 0.02861
0.15109 0.09275
0.06565 0.06686
0.38020* 0.35791**
0.166521 0.159569
0.33783* 0.31695**
0.22106* 0.15461
-0.00521 -0.00805
-0.05553 -0.02788
0.32147* 0.22396*
0.07986 0.07985
0.43089* 0.42012** 0.42751* 0.44681**
** and *: Significant at P < 0.01 and P < 0.05 respectively
Fertile nodes on secondary branch -0.38676* -0.37825**
Pod Length
Pod Breadth
Seeds pod-1
Pods plant-1
100 seed weight
Biomass plant-1
0.003372 0.00416
0.20656* 0.08513
-0.13114 -0.13018
-0.24761* -0.24573*
-0.16387 -0.16318
-0.38520* -0.38249**
0.27546* 0.26483**
0.07966 0.07975
0.07065 0.03863
0.03041 0.02974
0.25396* 0.24724*
0.06202 0.05963
0.20937* 0.20561*
0.28601* 0.27972**
-0.05575 -0.05541
0.22774* 0.16478
-0.07892 -0.06080
-0.15724 -0.05649
0.05429 0.03882
0.14942 0.10910
0.01655 0.01284
0.06254 0.06028
0.16401 0.13214
-0.12729 -0.08106
-0.02781 -0.02409
0.02799 0.02708
-0.30242* -0.29489**
-0.51854* -0.21967*
-0.02002 -0.01993
0.24719* 0.24372*
-0.21896* -0.21499*
0.08075 0.08058
0.17327 0.16953
-0.06813 -0.06411
0.46271* 0.42139**
0.30772* 0.28485**
-0.04610 -0.04615
0.05608 -0.00341
0.06521 0.06239
0.05210 0.04599
-0.14455 -0.13815
0.01008 0.00449
0.10050 0.09098
-0.15158 -0.14606
0.17574 0.16089
0.39565* 0.37780**
-0.39243* -0.38607**
-0.29173* -0.22933*
0.03612 0.03452
0.44461* 0.43359**
-0.29694* -0.29147**
0.23500* 0.22698*
0.12323 0.11687
0.24885* 0.24345*
0.51223* 0.53370**
0.03821 0.03627
-0.21204* -0.05256
-0.07998 -0.07561
0.06889 0.08982
-0.06923 -0.07554
0.02579 0.03797
0.14419 0.15277
-0.18689 -0.17310
-0.16151 -0.15786
-0.51922* -0.18538
0.06308 0.06183
0.28851* 0.29816**
-0.12650 -0.13046
0.21159* 0.21579*
0.17731 0.18355
0.16876 0.16657
0.76559* 0.30379**
-0.12673 -0.12552
-0.31280* -0.30914**
0.60092* 0.59712**
0.04510 0.04425
0.05833 0.05758
-0.01693 -0.01646
0.04457 -0.00133
-0.72757* -0.27890**
0.35771* 0.13781
-0.34704* -0.13173
-0.36589* -0.13702
-0.27474* -0.10416
-0.08405 -0.08405
0.23717* 0.23665*
0.17583 0.17539
0.21333* 0.21233*
-0.12673 -0.12979
0.74181* 0.73886**
0.64788* 0.64475**
0.48259* 0.47913**
0.43024* 0.42558**
0.35623* 0.35109**
0.31298* 0.31019**
0.83323* 0.83313**
0.70779* 0.70304**
0.09998 0.09923
Harvest Index plant-1 -0.11650 -0.11527
Seed yield plant-1 -0.52144* -0.51835**
0.23181* 0.22803*
Degree of Freedom: 90
50
index (rg = 0.232*, rp = 0.228*) and seeds per pod (rg = 0.213*, rp = 0.212*). Days to flower showed negative both phenotypic and genotypic correlations with seed yield per plant (rg = −0.521*, rp = −0.518**), and also with other yield related traits such as fertile nodes on primary (rg = −0.308*, rp = −0.302**) and secondary (rg = −0.387*, rp = −0.378**) branch, pods per plant (rg = −0.248*, rp = −0.246*) and biomass per plant (rg = −0.385*, rp = −0.382**). Plant height showed positive correlations with nodes on primary (rg = 0.380* and rp = 0.358**) and secondary (rg = 0.338* and rp = 0.317**) branch, fertile nodes on secondary branch (rg = 0.275* and rp = 0.265**), pods per plant (rg = 0.254* and rp = 0.247*), biomass per plant (rg = 0.209* and rp = 0.206*) and harvest index (rg = 0.286* and rp = 0.280**). Primary branches per plant had positive correlations with secondary branches (rg = 0.221*), nodes (rg = 0.321* and rp =0.224**) and fertile nodes (rg = 0.228*) on secondary branch. Secondary branches per plant showed positive correlations with fertile nodes on primary branch (rg = 0.431* and rp = 0.420**) and pods per plant (rg = 0.247* and rp = 0.244*) whereas with pod length (rg = −0.302* and rp = −0.295**) and breadth (rg = −0.519* and rp = −0.220*), and 100 seed weight (rg = −0.219* and rp = −0.215*), negative associations were observed. Nodes on primary branch showed positive correlations with fertile nodes on primary branch (rg = 0.428* and rp = 0.447**), and nodes (rg = 0.463* and rp = 0.421**) and fertile nodes (rg = 0.308* and rp = 0.285**) on secondary branch. Positive correlations of fertile nodes on primary branch were observed with fertile nodes on secondary branch (rg = 0.396* and rp = 0.378**), pods per plant (rg = 0.445* and rp = 0.434**) and biomass per plant (rg = 0.235* and rp = 0.227*) whereas negative correlations were observed with pod length (rg = −0.392* and rp = −0.386**), pod
51
breadth (rg = 0.292* and rp = 0.229*) and 100 seed weight (rg = −0.297* and rp = −0.292**). Nodes on secondary branch showed positive correlations with fertile nodes on secondary branch (rg = 0.512* and rp = 0.534**) whereas it showed negative correlation with pod breadth (rg = −0.212*). Fertile nodes on secondary branch showed positive correlations with pods per plant (rg = 0.289* and rp = 0.298**) and biomass per plant (rg = 0.212* and rp = 0.216*) whereas negative with pod breadth (rg = −0.519*). Pod length was positively correlated with pod breadth (rg = 0.766* and rp = 0.304**) and 100 seed weight (rg = 0.601* and rp = 0.597**) but it showed negative association with pods per plant (rg = 0.313* and rp = −0.309**). Pod breadth showed positive association with 100 seed weight (rg = 0.358*) but negative correlations were observed with pods per plant (rg = −0.728* and rp = −0.279**), biomass per plant (rg = −0.347*) and harvest index (rg = −0.366*). Seeds per pod correlated positively with biomass per plant (rg = 0.237* and rp = 0.237*). Pods per plant showed positive associations with biomass per plant (rg = 0.742* and rp = 0.739**) and harvest index (rg = 0.648* and rp = 0.645**). 100 seed weight positively correlated with biomass per plant (rg = 0.430* and rp = 0.426**) and harvest index (rg = 0.356* and rp = 0.351**). Biomass per plant showed positive association with harvest index (rg = 0.833* and rp = 0.833**). Path Coefficient Analysis: The genotypic correlations of various plant characteristics with seed yield per plant were further partitioned into direct and indirect effects. The pathways through which the yield components of lentil operate to produce their association with seed yield are depicted in table 3.3. Results showed that biomass per plant had the highest positive effect (1.437) on seed yield followed by pods per plant (0.249), pod breadth (0.186), fertile nodes on secondary branch (0.185), 100 seed weight (0.160) and secondary branches per plant (0.114).
52
Table 3.3: Direct and indirect effects of different characteristics on seed yield in lentil germplasm Parameter
Direct effect
Days to flower
Plant height
Primary branches plant-1
Secondary branches plant-1
Nodes on primary branch
Fertile nodes on primary branch
Days to flower
–0.024
1
0.02320
0.00491
–0.00290
–0.00135
0.04040
Plant height Primary branches
–0.133
0.00420
1
–0.01774
0.00747
–0.00972
Fertile nodes on secondary branch
Pod length
Pod breadth
Seed s pod-1
Pods plant-1
100 seed weight
Biomass plant-1
0.00004
–0.07141
–0.00060
0.03840
–0.00374
–0.06156
–0.02630
–0.55350
0.12800
–0.5214
–0.02187
0.00066
0.05086
–0.01294
0.01312
0.00867
0.06312
0.00994
0.30087
–0.31203
–0.0557
–0.117
–0.00102
–0.02004
1
0.02516
0.00013
0.00729
0.00063
0.04205
0.01282
0.02920
0.00155
0.03714
0.00265
0.08987
–0.17893
–0.1273
0.114
0.00061
–0.00871
–0.02595
1
–0.00204
–0.05658
–0.00005
0.00517
0.04914
–0.09631
–0.00057
0.06145
–0.03511
0.08987
–0.18903
–0.0681
–0.026
–0.00128
–0.05042
–0.00061
0.00909
1
–0.05614
0.00091
0.05682
0.00749
–0.01042
0.00178
0.01295
–0.02318
0.01449
–0.10964
–0.1516
–0.131
0.00743
–0.02208
0.00652
0.04905
–0.01092
1
0.00034
0.07306
–0.06374
0.05418
0.00103
0.11053
–0.04761
0.33771
–0.13444
0.2489
0.002
–0.00074
–0.04480
–0.03774
–0.00317
–0.01182
–0.02308
1
0.09458
–0.00621
–0.03938
–0.00282
0.01713
–0.01110
0.03706
–0.15731
–0.1869
0.185
–0.00935
–0.03653
–0.02674
0.00319
–0.00786
–0.05195
0.00100
1
0.02623
–0.09643
0.00180
0.07172
–0.02028
0.30406
–0.19340
0.1688
–0.162
–0.00009
–0.01056
0.00927
–0.03440
0.00118
0.05153
0.00008
–0.02982
1
0.14219
–0.00362
0.07775
0.09635
0.06481
–0.06364
–0.0170
Pod breadth
0.186
–0.00499
–0.00937
0.01846
–0.05903
–0.00143
0.03821
–0.00042
–0.09587
–0.12434
0.00127
–0.18087
0.05736
0.49871
0.39917
–0.2747
Seeds pod-1
0.029
0.00317
–0.00403
–0.00637
–0.00228
–0.00167
0.02058
0.0083
0.02486
0.02058
0.00828
1
0.02485
–0.01348
0.34082
–0.16182
0.2133
Pods plant-1
0.249
0.00598
–0.03368
–0.01754
0.02814
–0.00133
–0.05838
0.00014
0.05327
0.05080
–0.13513
0.00285
1
–0.02032
1.0660
–0.70681
0.4826
100 seed weight
0.160
0.00396
–0.00823
–0.00194
0.02493
0.00369
0.03899
–0.00014
–0.02326
–0.09760
0.06644
–0.00240
–0.03150
1
0.61827
–0.38860
0.3129
Biomass plant-1
1.437
0.00931
–0.02777
–0.00734
0.00919
–0.00026
–0.03086
0.03907
0.03907
–0.00732
–0.06445
0.00677
0.18441
0.06898
1
–0.90906
0.708
–1.091
0.00282
–0.03793
–0.01925
0.01972
–0.00257
–0.01618
0.00028
0.03274
–0.00947
–0.06795
0.00502
0.16106
0.05712
1
0.2318
plant-1
Nodes on secondary branch
Harvest index plant-1
Genotypic correlation (rg) with seed yield plant-1
Secondary branches plant-1
Nodes on primary branch Fertile nodes on primary branch Nodes on secondary branch Fertile nodes on secondary branch Pod length
Harvest index plant-1
53
1
1.1974
The positive indirect effect of biomass per plant on seed yield per plant via pods per plant (0.184) and 100 seed weight (0.069) was observed, followed by negative effect mainly via harvest index (−0.909). Indirect effect of the second most important character (pods per plant were) was observed mainly through biomass per plant (1.066) and fertile nodes on secondary branch (0.053) followed by negative effect of harvest index (−0.707). In the same way, pod breadth and fertile nodes on secondary branch also showed high positive direct effect on seed yield. Pod breadth showed positive effect (0.186) mainly via biomass (0.499) and harvest index per plant (0.399) on seed yield with negative effects via pods per plant (−0.181) and pod length (−0.124). Fertile nodes on secondary branch had high positive direct effect (0.185) . The indirect effect of fertile nodes on secondary branch were mainly through biomass (0.304) and pods (0.072) per plant followed by negative effects of harvest index (−0.193) and pod breadth (−0.096). Direct negative effects of harvest index (−1.091), pod length (−0.162), plant height (−0.133) and fertile nodes on primary branch (−0.131) followed by primary branches (−0.117) and days to flower (−0.024) showed adverse effects on seed yield. Harvest index showed the highest negative effect (−1.091) on seed yield manifested through combined negative effects of pod breadth (−0.068), plant height (−0.038), primary braches (−0.019) and fertile nodes on primary branch (−0.016) followed by nodes on primary branch (−0.003) and pod length (−0.010) whereas it showed positive correlation (0.232) with seed yield due to the high positive indirect effects through biomass (1.197) and pods (0.161) per plant. Indirect effect of pod length was observed via negative effects of mainly harvest index (−0.064), secondary branches (−0.034) and fertile nodes on secondary branch (−0.030). Indirect effect of plant height was manifested via harvest index (−0.312), fertile nodes on primary branch (−0.021), primary branches (−0.017) and pod length (−0.012). Fertile nodes on primary branch showed its indirect effect via
54
harvest index (−0.134), pod length (−0.063), 100 seed weight (−0.048) followed by plant height (−0.022) and nodes on primary branch (−0.011). Primary branches manifested their indirect effect mainly through harvest index (−0.179). Days to flower had indirect effect mainly through biomass (−0.554), fertile nodes on secondary branch (−0.071), and pods per plant (−0.062). Multiple Correlation and Regression Analysis: Detailed information on the effectiveness of different plant traits and their contribution toward final seed yield was estimated by multiple correlations (Table 3.4). This was accomplished by assessing the combined effect of yield components on seed yield keeping seed yield per plant as a dependent variable and other traits as independent variables. The relative contribution (R2) of 19.8 % toward seed yield was observed for days to flower, plant height and primary branches while the contributions of other trait combinations comprising of pod breadth, seeds per pod, and pods per plant; seeds per pod, pods per plant and 100 seed weight; pods per plant, 100 seed weight and biomass per plant; and biomass per plant, harvest index and days to flower were observed 58.6, 83.2, 83.6 and 95.1 % respectively. Contributions of all characters under study, yield related traits (pod length and breadth, seeds per pod, pods per plant, 100 seed weight, biomass per plant and harvest index) and other morphological characters (days to flower, plant height, primary and secondary branches, nodes on primary and secondary branches, and fertile nodes on primary and secondary branches) were observed 96.5, 95.0 and 21.9 % respectively. Other combinations of variables had less than 10% contribution (1.3-7.5 %) toward seed yield per plant. The joint association of all the characters with seed yield per plant through multiple correlations was highly significant (0.982**) followed by yield related traits (0.974**) and morphological characters (0.468*). Other combinations of variables pod breadth, seeds per pod, and pods per plant; seeds per pod, pods per plant and 100 seed
55
Table 3.4: Multiple correlation analysis of seed yield with different combinations of characters in lentil germplasm Parameter R R2 Adjusted S. E. 2 (%) R ** 0.445 ** 0.171 1.684 Y with X , X , X 19.8 1
2
3
Y with X2, X3, X4
0.229
5.2
0.020
1.831
Y with X3, X4, X5
0.115
1.3
−0.020
1.868
Y with X4, X5, X6
0.257
6.6
0.034
1.818
Y with X5, X6, X7
0.262
6.9
0.037
1.815
Y with X6, X7, X8
0.273
7.5
0.043
1.810
Y with X7, X8, X9
0.252
6.3
0.032
1.820
Y with X8, X9, X10
0.242
5.8
0.026
1.825
Y with X9, X10, X11
0.231
5.3
0.021
1.830
0.766 **
58.6 **
0.572
1.210
0.912 **
83.2 **
0.826
0.771
0.914 **
83.6 **
0.831
0.761
0.975 **
95.1 **
0.949
0.416
0.468 *
21.9 *
0.144
1.711
0.974 **
95.0 **
0.945
0.432
0.982 **
96.5 **
0.958
0.380
Y with X10, X11, X12 Y with X11, X12, X13 Y with X12, X13, X14 Y with X14, X15, X1 Y with Morphological characters (X1, X2, X3, X4 X5, X6, X7, X8) Y with Yield related traits (X9, X10, X11, X12, X13, X14, X15) Y with All characters Y X1 X4 X6 X8 X11 X15 S. E. ** & *
= Seed yield plant-1 = Days to flower (50%); X2 = Plant height; X3 = Primary branches plant-1 = Secondary branches plant-1; X5 = Nodes on primary branch = Fertile nodes on primary branch; X7 = Nodes on secondary branch = Fertile nodes on secondary branch; X9 = Pod length; X10 = Pod breadth = Seeds pod-1; X12 = Pods plant-1; X13 = 100 seed weight; X14 = Biomass plant-1 = Harvest index plant-1; R = Multiple correlation; R2 = Coefficient of determination = Standard error = Significant at P < 0.01 and P < 0.05 respectively
56
weight; pods per plant, 100 seed weight and biomass per plant; biomass per plant, harvest index and days to flower; days to flower, plant height and primary branches per plant showed positive associations with seed yield (0.766**, 0.912**, 0.914**, 0.975** and 0.445** respectively). Genetic Parameters: Plants within a random population vary in their expression of a particular quantitative trait due to genetic differences among the plants and to differences in the microclimate surrounding each plant, the variation described by phenotypic variance. Genotypic variance indicates the potential for the improvement of a particular trait. Heritability estimates enable the breeder in recognition of genetic differences among the traits and estimation of gain that may be accomplished by one generation of selection from the population being studied. Heritability in broad sense is the effectiveness with which the selection of genotype can be based on phenotypic performance. However, it has been suggested that heritability and genotypic coefficient of variation provide no indication for the amount of genetic progress that can be achieved through selection. But the estimates of heritability in conjunction with genetic advance indicate the feasibility of improvement in different traits. These estimates provide the existing stability, nature of variability and future selection possibilities in the breeding material. The results regarding estimates of genetic parameters are shown in table 3.5. The highest value of phenotypic coefficient of variation (PCV) was noted for pods per plant (27.57) followed by days to flower (2.28), harvest index (1.98) and seed yield (1.85) per plant. Other characters showed lower estimates of both PCV and GCV (genotypic coefficient of variation). Estimates of broad sense heritability among the traits under study ranged from 25.0 % (pod breadth) to 99.6 % (days to flower). High heritability estimates of biomass per plant (98.5 %), pods per plant (98.9 %), 100 seed weight (99.3 %), fertile nodes on secondary branch (95.9 %) and yield per plant (98.1 %)
57
Table 3.5: Estimates of phenotypic and genotypic variances, heritability and genetic advance for different plant characters in lentil germplasm Parameter Mean Phenotypic Genotypic Phenotypic Genotypic Heritability Genetic Variance Variance C. V. C.V. (h2) (%) Advance (broad sense) (% of mean) Days to Flower Plant height Primary branch plant-1 Secondary branch plant-1 Nodes on primary branch Fertile nodes on primary branch Nodes on secondary branch Fertile nodes on Secondary branch Pod Length Pod Breadth Seed pod-1 Pods plant-1 100 seed weight Biomass plant-1 Harvest Index plant-1 Seed yield plant-1
116.87±16.34 37.24± 6.12 2.63± 0.19 18.95± 4.91 23.41± 4.01 12.71± 3.78 19.01± 3.47 9.45± 3.44 1.18± 0.17 0.62± 0.09 1.62± 0.22 168.25±68.11 2.17± 0.84 14.69± 5.22 25.10± 7.05 3.82± 1.85
266.91 37.44 0.04 24.19 16.05 14.31 12.04 11.85 0.03 0.04 0.05 4638.63 0.70 3.39 49.76 27.24
265.84 35.25 0.02 23.26 15.02 13.91 11.04 11.37 0.03 0.01 0.05 4588.61 0.70 3.34 49.19 26.72
C.V.: Coefficient of Variation
58
2.28 1.01 0.01 1.28 0.69 1.13 0.63 1.25 0.02 0.06 0.03 27.57 0.32 0.89 1.98 1.85
2.27 0.95 0.07 1.23 0.64 1.09 0.58 1.20 0.02 0.01 0.03 27.27 0.32 0.88 1.96 1.82
99.6 94.2 50.0 96.2 93.6 97.2 91.7 95.9 99.2 25.0 99.4 98.9 99.3 98.5 98.9 98.1
28.68 31.87 7.22 51.48 32.98 59.56 34.46 72.06 29.66 9.38 27.78 82.49 79.26 98.94 56.51 71.82
were noted coupled with their respective high genetic advance expressed as percent of mean (98.9, 82.5, 79.3, 72.1 and 71.8 respectively). Principal Component Analysis: Since strong and very strong associations were observed among matrix correlations, it is needed to identify characters determining the improvement in lentil seed yield using principal component analysis (PCA). From PCA, first six principal components (PC1 – PC6) with eigenvalue > 1 were selected, and these components accounted for more than 75 % of the cumulative variance (Table 3.6). The remaining components were eliminated and considered less significant so that fewer components were dealt with. Based on the component loadings after the varimax rotation, the first eight components were extracted and the other components were eliminated. These first eight principal components accounted for 85.6 % of the total variance of the original data and the communalities showed that all the variables had been described to an acceptable level as communalities ranged from 0.75 to 0.973. The first component (PC1) gave information about the variation in pods, biomass and seed yield per plant which described more than 23 % of the variance. In this component, pods and biomass per plant were observed more important for the improvement of lentil seed yield. The second component described about 18 % variation, which originated mainly from pod/seed related traits. Nodes and fertile nodes on primary branch, and fertile nodes on secondary branch constituted about 13 % of total variance (PC3), days to flowering and harvest index described about 9 % of total variance (PC4), secondary branches per plant and fertile nodes on primary branch showed 7 % of total variance (PC5) and PC6 extracted more than 6 % of total variance described by primary branches per plant. First six PCs were observed
more
important
(75.7
%
of
59
the
cumulative
variance)
as
these
Table 3.6: Principal components analysis (PCA), varimax rotated factor loadings and communalities for variation among sixteen quantitative traits in lentil germplasm Parameter PC1 PC2 PC3 PC4 PC5 PC6 PC7 PC8 Eigenvalue % of variance explained % of cumulative variance explained
3.7269 23.3 23.3
2.8260 17.7 41.0
2.0667 12.9 53.9
1.3552 8.5 62.3
1.1135 7.0 69.3
1.0236 6.4 75.7
0.8509 5.3 81.0
Rotated loading
Days to flower Plant height Primary branches plant-1 Secondary branches plant-1 Nodes on primary branch Fertile nodes on primary branch Nodes on secondary branch Fertile nodes on secondary branch Pod length Pod breadth Seeds pod-1 Pods plant-1 100 seed weight Biomass plant-1 Harvest index plant-1 Seed yield plant-1
Communality
Factor 1
Factor 2
Factor 3
Factor 4
Factor 5
Factor 6
Factor 7
Factor 8
-0.098 0.169 0.037 0.127 -0.040 0.145 0.068 0.123 -0.053 -0.113 0.115 0.809 0.317 0.902 0.490 0.908
-0.034 0.041 -0.034 -0.197 0.027 -0.293 0.041 -0.175 0.912 0.859 -0.131 -0.330 0.750 0.115 0.096 0.137
-0.024 0.234 0.150 -0.133 0.659 0.385 0.848 0.731 -0.007 0.065 -0.030 0.124 -0.181 0.083 -0.076 -0.001
-0.907 0.079 -0.034 -0.051 -0.186 0.313 -0.120 0.430 -0.040 0.003 0.098 0.193 0.140 -0.096 0.673 0.337
-0.063 0.048 0.087 0.845 0.357 0.673 -0.072 -0.041 -0.156 -0.050 -0.054 0.162 -0.159 0.063 -0.009 0.041
-0.010 0.013 0.917 0.231 -0.260 -0.224 0.196 0.158 -0.011 -0.102 0.055 -0.004 0.090 0.121 -0.143 -0.039
0.166 -0.917 -0.013 -0.060 -0.264 0.020 -0.123 -0.064 -0.081 0.064 -0.001 -0.047 -0.042 -0.177 0.279 -0.026
0.044 -0.004 -0.055 0.081 -0.254 -0.018 0.105 0.045 0.025 0.115 -0.948 0.068 0.010 -0.100 -0.143 -0.110
60
0.7367 4.6 85.6
0. 868 0.935 0.878 0.853 0.800 0.857 0.810 0.798 0.869 0.786 0.945 0.850 0.750 0.904 0.827 0.973
principal components were related more to yield components, productive branches and phenology of the plants.
Discussion The information on variability of seed yield and its components is essential for the successful use of the material, and for defining selection criteria in the plant improvement programmes. In this study, results showed that the germplasm was amenable as a range of genetic variability was observed in all the characters showing highly significant differences. Greater variations were observed for nodes and fertile nodes on secondary branch, and seed yield per plant (Table 3.1). Various researchers (Balyan and Singh, 1986; Bicer and Saker, 2004a; Erskine, 1985; Erskine et al. 1990 and 1994b; Mia et al. 1986; Shahi et al. 1986; Singh and Rana, 1993; Stoilova and Pereira, 1999; Tullu et al. 2001; Zaman et al. 1989) have described wide ranges of variation for different phenological and morphological plant characteristics including seed yield per plant to identify the most promising genotypes for their use in the lentil improvement programmes, and breed higher biomass and seed yielding cultivars. Significant genetic variations for plant growth characters and specific adaptation have also been observed in germplasm collections for crop improvement (Cutforth et al. 2007). Photoperiod requirements of macrosperma type (exotic) lentils from latitudes farther from the equator have been described precluding their direct use (introduction) in this region (Erskine and Hawtin, 1983). In this study, the genotypes ILL 4679 (Argentina, early flowering), ILL 68 (Lebanon, late flowering), ILL 5748 (Syria, late flowering) and ILL 6024 (Syria, normal flowering), though performing well with respect to seed yield per plant under the prevailing weather conditions due to more rains (Table 2.2), may not be adapted in Pakistan due to their photoperiod sensitivity and inability to set pods at high temperatures and low humidity during the month of March (flowering 61
period and pod development stages) and onward (pod setting and seed development stages), thereby imbalancing the net photosynthetic rate during rapid seed development. The adverse effects associated with heat stress due to rise in temperatures and relatively low humidity (less than 50%) in lentil integrated with other physiological effects on pod/seed developments causing forced maturity, and ultimately leading to lower seed weight and seed yield have been described by several researchers (Chandra and Asthana, 1988; Erskine et al. 1994a; Muehlbauer et al. 2006; Summerfield, 1981; Summerfield et al. 1989). In spring wheat, higher air temperatures resulted in yield loss of 20% by reducing the maturity (Laurila, 2001). Temperature exceeding 30 to 32 oC has been observed a key limiting factor in chickpea yield by hastening maturity and/or by decreasing seeds per plant and seed weight (Harris, 1979; Wang et al. 2006), and increases in temperature above 2.5 oC would have overall negative effects on world agriculture (IPCC 2001; Smith and Almaraz, 2004). Cumulative rainfall and its distribution, and temperature fluctuations, particularly at flowering and pod setting stages, have been observed strong determinants of seed and straw yields. If more moisture is available to lentil even at near maturity stage, it continues new flower development and pod-filling, directly adding to seed yield and indirectly through a delay in maturity. But on the other hand, rise in temperature imposes negative effect on vegetative and reproductive growth phases and leads to forced maturity resulting decline in seed yield (Sarker et al. 2003). Also, negative effects of increasing temperature on crop production have been described by Philips et al. (1996) and Yu et al. (2002a) in semiarid climate conditions. The same trends of rainfall and temperature were observed during this study (Table 2.2) as more rains started at flowering and pod setting stages coupled with substantial rises in temperature during the month of March and onward. Conseqently, late flowering genotypes could not perform
62
well under these weather conditions. For example, a tall (43.6 cm plant height) and late flowering (129 days to 50% flowering) genotype ILL 2465 bearing higher number of pods (338) produced high biomass (25.4 g) could not cope with the increasing temperature with small seeds in pods (1.73 g per 100 seed weight) and low harvest index (27.6) impairing seed yield per plant (6.9 g) that might be attributable to short reproductive phase during pod/seed development. Likewise, Salam and Islam (1994) found that the production of higher biomass by lentil mutant MI 596 could not synchronize with high seed yield due to its inability of partitioning the assimilates towards pod development, and excessively delayed maturity under Indian conditions always lowered seed yield through reduced seed size (Tyagi and Sharma, 1985). Smaller seed size and seed growth rate at warmer environment have been explained as a result of smaller cell number without compensating increases in cell volume or cell growth rate in common bean (Sexton et al. 1997). While higher seed yields in lentil, chickpea and field pea have been realized under combination of high growing season rainfall and lack of excessively high temperatures during the flowering and pod-filling periods (Nielsen, 2001). He also observed that higher temperatures resulted in flower and pod abortion, and shortening of the seed-filling period culminated to lower seed yields which confirms the positive association of seed-filling duration with seed yield in lentil (Khan et al. 1998). Various physiological bases, causing seed yield to be low, have been suggested for lentil yield improvements in different studies. The associations of total dry matter, crop growth rate, net assimilation rate, relative growth rate, soluble protein content and harvest index with seed yield showed indications of prime importance of photosynthetic potentials of varieties and proper distribution of assimilates towards the reproductive sink for the maximal yield expression (Dutta and Mondal, 1998). High proportion of
63
unfilled pods in the upper part of the plant canopy has been explained due to the lack of assimilates to young pods developing during the late stages of the reproductive phase (Penaloza, 1985). Whitehead et al. (2000) suggested that improved seed yield was possible from increased vegetative biomass prior to flowering provided N-assimilation increased. The increased biomass would later be remobilized and translocated to developing seeds. Tullu et al. (2001) emphasized on the rapid crop growth, and higher dry matter partitioning to seed combined with early flowering and maturity, needed to be considered in the selection process. Associations of high seed yield with rapid ground cover, early flowering and maturity, long reproductive period, greater number of seeds and pods, high total dry matter, greater harvest index, and high water use efficiency have been observed in lentil (Shrestha et al. 2005). McPhee and Muehlbauer (2001) suggested that partitioning of photosynthates during the reproductive period in peas would be particularly important. As peas are indeterminate and continue to grow vegetatively when experienced to adequate moisture. Greater seed yields coupled with increases in seed size would result if cultivars could be developed which partition greater amounts of photosynthates to the seed late in the season, and Berger (2006) noticed that early phenology was required to avoid terminal drought and heat stress in chickpea. From this study, it was observed that most of the genotypes were late in flowering and could not cope with the rising temperatures to seed yield during prevailing growing conditions indicating the importance of phenology of the genotype. Low seed yields of grain legume species have been described attributable to later flowering and podding in southwestern Australia (Thomson et al. 1997) while on the other hand, the genotypes which were well-adapted to their environment maximized effective crop duration, resource capture and thereby dry matter accumulation, and ultimately biomass and seed yield (Whitehead et al. 1998). Thus, it seems important to follow an appropriate
64
procedure during selection through the incorporation of favouable plant characters in to the breeding materials rather than to direct selection (introduction). Researchers have emphasized to follow different strategies according to the prevailing environments for realization of better seed yields in lentil and in other crops. Hence, an understanding of the adaptation of the crop is basic to its effective improvement (Ceccarelli et al. 1994). Cutforth et al. (2007) suggested that developing new cultivars would require identification of crop traits that would allow the crop to respond well in the future climate, and the crops best adapted to the prevailing environmental factors tended to be the most productive (Miller et al. 2002). Therefore, selection of genotypes with early seedling establishment, early and rapid biomass development, and early flowering and maturity in sites of extremely low rainfall has been advocated (Andrews and McKenzie, 2007). Strong positive associations of seed yield per plant with fertile nodes on primary branch, seeds per pod, pods per plant, 100 seed weight, biomass per plant, and harvest index indicated the importance of these traits for selection of high-yielding genotypes under the prevailing weather conditions, provided the genotypes had appropriate phenology (negative association of days to flower with seed yield per plant). Direct roles of yield components on yield improvements have been observed in soybean and harvest index was observed the largest contributor to yield improvements compared with biomass (Cui and Yu, 2005). A number of reports are available on positive relationships of seed yield with seeds per pod, pods per plant, 100 seed weight, seed size, biomass and harvest index, and also with other plant traits (Baidya et al. 1988; Balyan and Singh, 1986; Chakraborty and Haque, 2000; Chauhan and Sinha, 1982; Chauhan and Singh, 2001; Naji et al. 2003; Ramgiry et al. 1989; Rao and Yadav, 1988; Singh and Gupta, 2005; Singh et al. 1989; Singh et al. 2003; Sinha and Singh, 2002; Tambal et al. 2000; Tyagi and
65
Sharma, 1985; Verma et al. 2004; Vir et al. 2001; Whitehead et al. 1998 and 2000). The phenology have been observed to be the most important single factor influencing crop adaptation since it largely determines when annual cereal, pulse and oilseed crops will subsequently ripe for harvest (Summerfield et al. 1997). In the present study, the strong positive association of seed yield per plant with fertile nodes on primary branch, and other yield related traits such as seeds per pod, pods per plant, 100 seed weight, biomass and harvest index might be attributed to the excessive rainfall during the months of February and March (vegetative and flowering stages), and the late flowering genotypes continued to grow and flowering, favouring more pods per plant but with unmature seeds in the pods (negative association of pods per plant with 100 seed weight) that might occurred due to afterward high temperatures during seed development stage. Decline in pod set with the rise in temperature has been observed in lentil (Chandra and Asthana, 1988), and proposed to use locally adapted germplasm and select in the target environment for productivity improvements of the crops (Ceccarelli et al. 1998) or to identify cultivars which had increased heat/water stress tolerance (Cutforth et al. 2007). Negative correlations of days to flower with seed yield per plant, and also with fertile nodes on primary and secondary branch, pods per plant, and biomass per plant, showed that phenology had significant effects in determining the seed yield of individual genotype. In the literature, reports indicated positive association of days to flowering with seed yield per plant but Kumar and Sapra (1984), Tyagi and Sharma (1985), Mia et al. (1986), Khan et al. (1998) and Kumar et al. (1995) observed negative relationship of flowering with seed yield. Rathi (2004) also found negative association of flowering with yield and pods per cluster. This study indicated that very late flowering genotypes continued to grow vegetatively producing less number of fertile nodes, low number of pods with filled seeds on these branches
66
causing low harvest index (negative association of days to flower with harvest index) and ultimately, less seed yield. It means that selection of genotypes possessing appropriate phenology with determinate growth habit is likely to produce better seed yields. The associations of plant height with nodes on primary and secondary branch, fertile nodes on secondary branch, pods per plant, biomass and harvest index, and relationships of pods per plant with morphological and other yield related traits indicated that these associations would favour the selection of tall bushy plants bearing more productive branches and produce more seed yield provided the phenology of the plants synchronized with the prevailing environments. Selection of profusely branched plants bearing high pod number has been suggested to produce higher yields (Zaman et al. 1989). However, large plant framework alone might not increase the efficiency of photosynthetic partitioning or improve seed yield (Silim et al. 1993). Normally, bushy plants with more primary and secondary branches have been observed to produce pods only on the periphery and the central portion remained barren. Resultantly, bushy plants produce relatively more plant dry matter than grain yield and lower harvest index (Singh, 1977). Other researchers (Erskine, 1996; Hamdi et al. 1991; Kusmenoglu and Muehlbauer, 1998) attempted to select indirectly for seed yield by selecting for positively associated characteristics, and larger vegetative frame was correlated positively with both seed and straw yields due to greater number of reproductive nodes and pods on taller and more branched plants. Thus, selection of tall bushy plants, with early vigour and more productive branches resulting in larger vegetative biomass, may likely favour to increased radiation interception and thereby optimizing seed yield. The attainment of high leaf area index, that reduced soil water evaporation, intercepted and converted radiation into dry matter efficiently, and partitioning of the dry matter to the seeds, has been described as the major requirement for high seed yield in grain legumes
67
in the semi-arid environments (Tesfaye et al. 2006), and during selection the resilience of pulse crops to present-day weather extremes such as excess water, heat, drought and cool weather during grain-filling phase may be considered to predict adaptation to future changes (Cutforth et al. 2007). Biomass per plant showed the highest direct positive effect on seed yield per plant followed by pods per plant, pod breadth, fertile nodes on secondary branch, 100 seed weight and secondary branches per plant indicating the importance of these characters during selection. The results of Singh and Gupta (2005) revealed that biological yield in lentil had the maximum direct effect on seed yield which confirmed the present findings. Positive indirect effects of biomass per plant, pods per plant, fertile nodes on secondary branch, 100 seed weight and secondary branches per plant were observed on seed yield per plant via pods per plant and 100 seed weight, biomass per plant and fertile nodes on secondary branch, biomass and pods per plant, biomass and pod breadth, and biomass and pods per plant respectively. Several scientific reports (Balyan and Singh, 1986; Chakraborty and Haque, 2000; Chauhan and Sinha, 1982; Dixit and Dubey, 1984; Nandan and Pandya, 1980; Singh, 1977) indicated large direct effects of number of pods, seeds per pod, secondary branches and seed size on grain yield, and very strong influences of biological yield and pods per plant on grain yield have been described in lentil (Balyan and Singh, 1986; Kumar et al. 2004b; Luthra and Sharma, 1990). Present studies showed that biomass and pods per plant may contribute to improved seed yield per plant directly. It means that genotypes having larger biomass with higher number of pods may be identified during selection. The positive direct effects of biomass per plant in this study may be attributed due to rains at the flowering time as most of the genotypes were late in flowering. The positive effect of increased soil moisture on biological yield of lentil has been observed when irrigated (Ibrahim et al. 1993).
68
Harvest index showed the highest negative direct effect on seed yield and its indirect influence manifested mainly via plant height, primary branches and fertile nodes on primary branch. The excessive amount of rainfall apparently allowed plants to grow vegetatively after entering the reproductive phase. As a consequence of high biomass production, a negative direct effect of harvest index was observed on seed yield attriburable due to the effect of high temperatures during pod setting stage, thus producing immature seeds. A positive influence of harvest index might be occurred if the plants under mild temperatures had a longer season to produce additional seeds. But in our conditions, high temperatures during reproductive phase do not allow plants to develop pods with mature seeds resulting in low harvest index and ultimately lower seed yields. Seed yield might be increased selecting large vegetative biomass while keeping the harvest index constant (Kusmenoglu and Muehlbauer, 1998). Increased seed yield in lentil has been observed, almost entirely accounted for by increased harvest index (Hawtin et al. 1980) while a linear and positive relationship between biomass and seed yield has been observed in common bean (Scully and Wallace, 1990). The negative direct effects of harvest index may be overcome by applying appropriate selection criteria in a large population of diverse type of genotypes. This may be achieved either through selection of genotypes which establish greater vegetative growth before flowering (early vigour) or following partitioning of the dry matter in seed production. Such type of genotypes may utilize stored soil moisture and escape terminal drought and heat stress by maturing early. Nevertheless, consistent seed yield improvements have not been associated with consistent increases in harvest index. Environment and phenology play a large role in determining the harvest index of individual genotype. Biomass and seed yield may be increased further by the selection of cultivars in more variable populations developed through hybridization (Whitehead et al. 2000). The more early growth rate requires
69
greater amounts of water and nutrition to convert the light energy into carbohydrates or through early flowering genotypes with late maturity ultimately gaining longer reproductive period (grain-filling). This could be possible by selecting higher biomass producing genotypes with equivalent or higher harvest index. Low harvest index may also be attributed in this study due to inefficient resource capture by the late flowering genotypes that reduced seed yield (negative association between harvest index and days to flower). Low crop harvest indices in wheat have been speculated due to poor conservation of light, water and nutrients to grain, indicating that potential yields were not being realized (Riffkin and McNeil, 2006). The negative effects of harvest index may be nullified by selecting more determinate genotypes in which flowering period is condensed and a greater proportion of nodes bear pods. In such type of genotypes, assimilates may be partitioned to pods rather than to continued vegetative growth, and the production of late pods, which do not produce mature seeds by the time of harvest. Multiple correlation studies showed that yield related characters (pod length and breadth, seeds per pod, pods per plant, 100 seed weight, biomass per plant and harvest index) along with days to flower were the major components determining seed yield. Major changes in lentil seed yield have been observed relating to the characters such as pods, seed weight and size of leaflet (Safaei, 2001) while the contributions of plant height, branches per plant and pods per plant have also been observed better than other characters (Jain et al. 1991). In the present studies, morphological characters had less effects on seed yield compared with yield related traits. The same trend of these morphological characters was confirmed in path coefficient analysis studies. However, these traits showed adverse indirect effects on seed yield per plant via other yield related traits. For example, days to flower influenced seed yield indirectly via pods per plant, 100 seed weight and biomass per plant. Likewise, plant height contributed detrimental indirect effects on seed yield via
70
pod length and breadth, seeds per pod, pods per plant, seed weight, biomass and harvest index. Primary branches influenced seed yield via harvest index. Nodes and fertile nodes on primary and secondary branches affected seed yield per plant indirectly through harvest index. From the relationships of seed yield with other component characters and relationships within component characters suggested that selection criteria should be formulated based on component characters comprising of morpho-phenological traits especially days to flowering along with yield relating traits for exploitation of improved seed yield. Plant height has been suggested to be used for indirect selection for increased biomass, and seed yield coupled with constant harvest index (Kusmenoglu and Muehlbauer, 1998). Days to flower (phenology), productive branches and yield related traits seemed to be given due consideration during selection for realizing potential seed yields. Higher values of phenotypic coefficients of variance (PCV) than those of genotypic coefficients of variance (GCV) indicated the environmental influence upon the characters studied that is supportive to findings of Chauhan and Singh (1998) in the improvement of lentil. The highest values of PCV and GCV observed for pods per plant followed by days to flower indicated the more effect of environmental conditions on these traits. Singh et al. (1995), Vir et al. (1995 and 1998) and Safaei (2001) also observed high values of PCV and GCV for pods per plant as well as for seed yield, biomass, harvest index and seed weight. High heritability estimates coupled with high genetic advance of biomass per plant, pods per plant, 100 seed weight, fertile nodes on secondary branch and seed yield per plant followed by moderate estimates of fertile nodes on primary branch, harvest index and secondary branches in this study indicated that improvements in these characters might be possible within the examined genetic material keeping phenology of the genotypes suited to the prevailing environments as
71
days to flower (high heritability with low genetic advance) was observed the second most influencing character in this study. High heritability estimates for days to flowering with low genetic advance has been observed in lentil (Baidya et al. 1988). Characters possessing high heritability (h2) estimates combined with higher genetic advance (GA) are controlled by additive type of genes and may be used directly for the improvement of seed yield whereas the characters having high h2 along with low GA are controlled by non-additive type of genes (dominant, epistasis or their interactions), for which appropriate selection strategies have been suggested (Johnson, 1955; Sarwar et al. 2004). If the variation is controlled by additive effects and the quantitative trait has intermediate heritability, then this approach might be quite powerful (Hoffmann and Willi, 2008). Several scientists (Chakraborty and Haque, 2000; Daychand, 2007; Jain et al. 1995; Ramgiry et al. 1989; Rathi et al. 2002; Singh and Singh, 1969) observed high estimates of heritability in conjunction with high genetic advance for days to maturity, number of branches per plant, seeds per pod, 100 grain weight (seed size), pods per plant, seed yield per plant and harvest index. Likewise, other researchers (Islam and Shaikh, 1978; Rahman and Sarwar, 1982; Saraf et al. 1985; Rajput and Sarwar, 1989; Ramgiry et al. 1989; Zaman et al. 1989) reported the greater importance of pod number as well as primary and secondary branches to seed yield. The harvest index is the most important trait in enhancing lentil production. But the relatively low h2 and GA estimates of most of the traits relating to harvest index such as plant height, primary and secondary branches, nodes and fertile nodes on primary branch, nodes on secondary branch, and seeds per pod further indicate that improvements in harvest index through breeding would be a very demanding task. In this study, higher estimates of heritability for pods per plant and its strong positive association with plant height, secondary branches, and fertile nodes on primary and secondary branches, and negative correlations with days to
72
flower indicate the importance of phenology and tall bushy plants bearing more productive branches in integrating genotypic variation for these characters toward selection in lentil. There seems a potential to increase seed yield by selecting genotypes for greater early vigour in a lentil breeding programme. More than 70 % of the gross phenotypic diversity was explained on the basis of first six PCs in the principal component analysis (PCA) whike Sonnante and Pignone (2007) observed 53.3 % of the total variation accounted for by the first three PCs in lentil that is very similar to the present findings (53.9 %) and more than 23 % of the variance extracted by the first PC was due mainly to variations in the pods, biomass and seed yield per plant. Variations in pod length and breadth, and 100 seed weight contributed more than 15 % of the whole variance accounted for by the second PC. La`zaro et al. (2001) observed that seed weight, testa pattern and testa pattern colour in lentil described 32.4 % of the total variance accounted for by the first PC, plant height and first pod height showed 15.1 % of the total variance extracted by second PC and third PC extracted 10.7 % of the total variance explained by days to flowering and flowering duration while in this study, variations in days to flowering and plant height described 11.3 and 7.1 % of the total variance defined by fourth and seventh PC respectively that might be attributed due to the differences in the materials studied. All of the traits under study appeared to have high contributions towards the gross phenotypic variability apparent among the genotypes, and it is possible to decide the importance of traits in different principal components (Johnson and Wichern, 2002). In this study, morphophenologic and yield related traits particularly productive branches, pods per plant, biomass per plant, harvest index and days to flowering seemed to have an ample role in the improvement of lentil seed yield.
73
Summarizing the results, it may be concluded that the population is amenable to selection for realizing potential seed yield under prevailing weather environments as significant variations were observed among the genotypes for all the characters under study. Flowering time seemed to have worth measuring in improving seed yield as substantial rise in temperatures during the month of March and onward (Table 2.2) cause low pod setting and low seed weight through forced maturity in the late maturing genotypes resulting in poor seed yield of individual genotype. Improvements in seed yield per plant may likely be favourable by selecting big plants possessing early vigour with more productive branches if the plants could partition greater amounts of photosynthates to the seeds rather than to additional vegetative growth and the production of late pods which do not produce mature seeds by the time of harvest. Strong positive associations of pods with seed yield as well as with secondary branches, fertile nodes on primary and secondary branches, and contrasting correlations with days to flower indicated the importance of phenology and bushy (more branched) plant type in integrating genotypic variation for these characters towards selection with improved seed yield under the prevailing weather conditions. However, the expression of productivity under field conditions and the translation of this improvement to higher seed yields depend on the environment and phenology, and the canopy structure of the crop (Hamid et al. 1990). The canopy structure of the crop in community is modified by different growing conditions which in turn regulates the efficiency of light utilization in crop productivity (Gifford et al. 1984). Alternatively, improvements may be realized through the selection of more determinate type of genotypes which establish greater vegetative growth before flowering (early vigour) followed by partitioning the dry matter to seed. Such genotypes may utilize stored soil moisture and escape terminal drought and heat stress by maturing early.
74
Chapter 4
Genetic Diversity in Parental Lines For the success of any hybridization programme, the choice of parents is of paramount importance. Diverse parents are expected to yield the best recombinants and a quantitative estimation of genetic diversity in any gene pool guides the breeder to make crosses between desirable but diverse genotypes to generate material for a selection programme (Gupta et al. 1996). Studies on genetic diversity provide important baseline for future germplasm collection/conservation and improvement programmes (Fikiru et al. 2007). Genetic diversity has been explored in cultivated and wild lentils by using morphological traits (Ahmad et al. 1997a) as well as RAPD markers (Abo-Elwafa et al. 1995; Sharma et al. 1995; Ahmad et al. 1996; Ferguson et al. 1998a and b; Duran et al. 2004). The breeder can use the genetic similarity information to complement phenotypic information in the development of breeding populations (Smith et al. 1990; Nienhuis et al. 1993; Galven et al. 2001; Pengelly and Lui, 2001; Greene et al. 2004). In recent years, molecular markers have been developed based on the more detailed knowledge of genome structure and considerable emphasis has been laid on the use of molecular markers in practical breeding and genotype identification (Ovesna et al. 2002). Different types of molecular markers have been used to assess the genetic diversity in crop species, but no single technique is universally ideal. The choice of the technique depends on the objective of the study, sensitivity level of the marker system, financial constraints, skills and facilities available (Yoseph et al. 2005). Among the molecular markers, RAPDs are particularly useful for the assessment of genetic diversity because of their simplicity,
75
speed and relatively low cost (Nybom, 2004) and can be used to characterize DNA variation patterns within species and among closely related taxa. Despite the limitations associated with randomly amplified polymorphic DNA (RAPD) markers, these markers have been widely used for the determination of genetic diversity, phenetic relationships, and the identification of cultivars and parents in a number of plant species (Ferguson et al. 1998c; Fernandez et al. 2002; Haley et al. 1994; Hou et al. 2005; Lee, 1995; Rao et al. 2007; Samec and Nasinec, 1995; Sonnate and Pignone, 2001; Tingey and del Tufo, 1993; Yüzbaşıoğlu et al. 2006). Nagella et al. (2008) reported that RAPD markers had a potential for identification and maintenance of niger germplasm for crop improvement purposes. The genetic diversity in the parental lines used for hybridization programme during the present studies was determined using morphological plant characteristics as well as RAPD markers.
Genetic diversity in parental lines using morphological characteristics The mean square, range and mean values for each character in parental lines are presented in table 4.1. Highly significant differences were observed among the parentals for all the characters under study. Coefficient of variation (C. V.) indicated that days to flower (0.99 %), 100 seed weight (1.75 %), plant height (2.56 %), harvest index (2.83%), primary branches (4.48 %) pods per plant (5.35 %), fertile nodes on primary branch (6.26 %), biomass (6.36 %) and seed yield per plant (6.68 %) had low to moderate variability. Greater variations were observed for fertile nodes on secondary branch (7.60 %) and secondary branches (7.99 %) in the parents. Flowering among the genotypes ranged from 65.5 (ILL 6821) to 100.3 days (ILL
76
Table 4.1: Values of means, coefficient of variance, mean square, and range for different plant characteristics in lentil parental lines Primary branches plant-1
Secondary branches plant-1
Fertile nodes on primary branch
Fertile nodes on secondary branch
Pods plant-1
100 seed weight (g)
Biomass plant-1 (g)
Harvest index plant-1 (%)
Seed yield plant-1 (g)
36.7c 39.9b 29.0f 44.6a 25.9g 36.9c 29.4f 22.8h 37.2c 33.6d 26.1g 31.5e
2.7bcd 3.1a 2.6cd 2.7bcd 2.9b 2.5d 2.2e 2.6bcd 2.7bc 2.7bc 2.6cd 2.6cd
10.7d 19.7a 12.8c 19.4a 14.0c 16.3b 9.3d 9.8d 16.1b 12.5c 5.9e 16.0b
9.4g 13.7bc 10.3fg 15.9a 12.9cd 14.4ab 10.9ef 9.5g 13.6bc 12.3de 6.5h 11.7de
6.8ef 9.1bc 6.5ef 11.9a 9.8b 11.4a 5.8f 7.2e 12.3a 8.5cd 4.3g 7.7de
151.4f 267.3a 134.6f 219.4c 180.6de 236.1b 106.5g 169.6e 188.6d 146.5f 135.4f 145.1f
3.4b 1.9i 2.6f 2.7e 2.4gh 1.8j 3.0c 2.3h 2.4g 2.9d 3.9a 1.9ij
16.3bc 20.5a 10.9g 13.0ef 13.9de 17.5b 14.9cd 16.3bc 11.5fg 12.9ef 12.9ef 12.2fg
34.0def 36.9c 32.1g 38.4bc 33.2efg 38.8b 30.0h 31.9g 35.2d 42.6a 32.4fg 34.4de
5.5c 7.6a 3.5g 4.9cde 4.6def 6.8b 4.5ef 5.2cd 4.1fg 5.6c 4.2f 4.2f
32.80 2.56 125.64**
2.65 4.48 0.14**
13.54 7.99 52.83**
11.72 6.26 18.99**
8.44 7.60 19.43**
152.06 5.35 6705.23**
2.60 1.75 1.19**
12.03 6.36 23.31**
34.89 2.83 38.43**
4.33 6.68 4.13
2.18 3.08
5.88 19.71
6.53 15.85
106.62 267.32
1.83 3.94
10.94 20.52
30.04 42.58
3.51 7.56
Parameter
Days to flower (50%)
Plant height (cm)
ILL 5782 ILL 2580 ILL 6024 ILL 8117 ILL 6468 ILL 7556 ILL 6821 ILL 7715 Masoor 93 Turk Masoor ILL 4605 Pant L 406
97.3c 98.6bc 98.1bc 97.3c 99.0ab 100.3a 65.5g 93.3d 97.0c 79.1f 80.9e 96.9c
Overall Mean 91.96 C. V. (%) 0.99 Mean Square 353.17** Range Low 65.53 High 100.33
22.83 44.55
Superscripts common in columns do not differ significantly
4.25 12.33
C. V.: Coefficient of variance
77
**: Significant at P < 0.01
7556). Plant height ranged from 22.8 (ILL 7715) to 44.6 cm (ILL 8117). Primary branches ranged from 2.2 (ILL 6821) to 3.1 (ILL 2580). Genotype ILL 2580 produced maximum secondary branches (19.7) while minimum (5.9) were noted in ILL 4605. Genotype ILL 8117 produced maximum fertile nodes (15.9) on primary branch and Masoor 93 on secondary branch (12.3) whereas ILL 4605 produced only 6.5 fertile nodes on primary branch and 4.3 on secondary branch. The genotype ILL 2580 produced maximum number of pods (267.3) whereas minimum (106.5) were recorded in ILL 6821. The largest seed size was observed in ILL 5748 (3.9 g per 100 seed weight) while ILL 7556 had minimum seed weight of 1.8 g. The genotype ILL 2580 produced the highest biomass per plant (20.5 g) as compared with the lowest biomass of 10.9 g (ILL 6024). The highest harvest index (42.6 %) was recorded in Turk Masoor whereas ILL 6821 had the lowest harvest index (30.0 %). Seed yield per plant among the genotypes ranged from 3.5 g (ILL 6024) to 7.6 g (ILL 2580). From principal component analysis (PCA), first three principal components (PC1 – PC3) with eigenvalue > 1 were selected (Table 4.2), and these components accounted for more than 80 % of the cumulative variance. The remaining components were eliminated and considered less significant so that fewer components were dealt with. Based on the component loadings after the varimax rotation, the first four factors were extracted and the other components were eliminated. These first four factors accounted for about 90 % of the total variance of the original data and the communalities showed that all the variables had been described to an acceptable level as communalities ranged from 79.2 to 98.5. The first component (PC1) gave information about the variation in plant height and harvest index which described more than 55 % of the total variance. In this component, big plants having more productive branches with high harvest index were
78
Table 4.2: Principal components (PCA) analysis, varimax rotated factor loadings and communalities for eleven quantitative traits in lentil parental lines Parameter PC1 PC2 PC3 PC4 Eigenvalue
6.100
1.709
1.191
0.873
Proportion
0.555
0.155
0.108
0.079
Cumulative
0.555
0.710
0.818
0.898
Rotated loading
Communality
Factor
Factor
Factor
Factor
1
2
3
4
Days to flower
0.050
0.544
0.008
0.743
0.850
Plant height
0.839
0.223
0.166
0.103
0.792
Primary branches plant-1
0.249
0.057
0.220
0.884
0.895
Secondary branches plant-1
0.570
0.698
0.118
0.327
0.933
Fertile nodes on primary branch
0.668
0.704
0.089
0.039
0.952
Fertile nodes on secondary branch
0.640
0.653
-0.028
0.206
0.879
Pods plant-1
0.436
0.483
0.533
0.455
0.914
100 seed weight
-0.047
-0.901
-0.225
-0.165
0.891
Biomass plant-1
-0.021
0.098
0.983
0.075
0.982
Harvest index plant-1
0.857
0.041
0.167
0.187
0.799
Seed yield plant-1
0.351
0.114
0.909
0.150
0.985
79
observed more important. 100 seed weight contrasted with fertile nodes on primary branch described by second component and it accounted for about 16% of the total variation. The third component explained nearly 11 % of the total variance constituted by biomass and seed yield per plant. Days to flowering and primary branches per plant contributed about 8 % of the total variance described by PC4. First four PCs were observed more important (89.8 % of the cumulative variance) as these principal components were related to yield contributing characters such as biomass, harvest index, productive branches, plant height and days to flowering. The cluster analysis resulted in the grouping of 12 genotypes into 4 major clusters comprising 2-5 genotypes (Fig. 4.1). In cluster A, three genotypes (Turk Masoor and Masoor 93 belonging to Pakistan, and ILL 8117 from ICARDA, Syria) were grouped while in cluster B, two genotypes from India (ILL 7556 and ILL 2580) were grouped. Two early flowering genotypes (ILL 4605 from Argentina and ILL 6821 from ICARDA, Syria) were grouped in cluster C. It is mentioned that one of the parents used in the development of ILL 6821 was ILL 4605. Cluster D consisted of a mixture of five genotypes belonging to India (ILL 7715 and Pant L 406) and ICARDA, Syria (ILL 5782, ILL 6024 and ILL 6468).
Genetic diversity in parental lines using randomly amplified polymorphic DNA (RAPD) markers The amplification profiles of twelve lentil genotypes (five macrosperma and seven microsperma types) with fifteen primers were polymorphic. There was not a single primer that could differentiate clearly among all the genotypes. The levels of polymorphism were different with different primers among different genotypes
80
81
(Fig. 4.2a, b and c). The number and size of amplified fragments varied with different primers. Maximum number of 9 fragments was amplified with primer OPN-13 while the primer OPB-9 produced only three fragments. The similarity matrices are shown in table 4.3. Data revealed that the similarity ranged between 87.2 % and 97.8 % among all the genotypes. From the similarity matrix, the least similarity was shown by the genotype ILL 6024 (red cotyledon and plain/large seeded genotype belonging to ICARDA, Syria). Molecular variability showed loose geographical parallelism as well as type of the genotype (macrosperma and microsperma type) in the clustering pattern (Fig 4.3). In the dendrogram, ILL 6024 (macrosperma type) did not cluster with any of the genotypes tested and was easily distinguishable. This genotype was crossed with ILL 8117 (in subcluster c, microsperma type). 88.3 % similarity matrix was observed between these genotypes. Rests of the genotypes were grouped in three main clusters. In cluster A, only two genotypes (ILL 2580 and ILL 7715) were grouped and their similarity coefficient was 90.3%. Both these genotypes (red cotyledon colour with spotted/small seeds) belonged to India. Genotype ILL 2580 (in cluster A, red cotyledon and spotted/smallseeded genotype belonging to India) crossed with ILL 5782 (in cluster C, red cotyledon and plain/large-seeds genotype belonging to ICARDA, Syria) were observed distinct and their similarity matrix was 89.2%. The other genotype ILL 7715 was crossed with ILL6821 (sub-cluster a, early flowering (66 days), quick growing habit, large seeds.) In cluster B, the genotypes belonging to ICARDA (Syria), India and Pakistan were grouped. This cluster can be divided in to three sub-clusters, each consisting of three, one, and two genotypes. In sub-cluster a, three genotypes belonging to Pakistan (Masoor 93 and Turk Masoor) and ICARDA, Syria (ILL 6821) had 97.8% similarity matrix among them whilst all the genotypes had contrasting morphological, growth habit and seed related traits. Genotype ILL 6821 (in sub-cluster a, from ICARDA, Syria) is early
82
Fig 4.2(a): Amplification of RAPD marker OPN-06 for lentil parental lines (1. ILL 4605, 2. ILL 5782, 3. ILL 6024, 4. Pant L 406, 5 ILL 8117, 6. ILL 7556, 7. ILL 7715, 8. ILL 6468, 9. ILL2580, 10. Masoor 93, 11. Turk Masoor, 12. ILL 6821)
1
2
3
4
5
6
7
8
9
10
11
12
Fig 4.2(b): Amplification of RAPD marker OPB-1 for lentil parental lines (1. ILL 4605, 2. ILL 5782, 3. ILL 6024, 4. Pant L 406, 5 ILL 8117, 6. ILL 7556, 7. ILL 7715, 8. ILL 6468, 9. ILL2580, 10. Masoor 93, 11. Turk Masoor, 12. ILL 6821)
Fig 4.2(c): Amplification of RAPD marker OPA-13 for lentil parental lines (1. LL 4605, 2. ILL5782, 3. ILL 6024, 4. Pant L 406, 5 ILL 8117, 6. ILL 7556, 7. ILL 7715, 8. ILL 6468, 9. ILL2580, 10. Masoor 93, 11. Turk Masoor, 12. ILL 6821)
83
Table 4.3: Estimates of similarity matrices in the parental lines using RAPD markers Genotype
ILL 4605 ILL 5782 ILL 6024 Pant L 406 ILL 8117 ILL 7556 ILL 7715 ILL 6468 ILL 2580 Masoor 93 Turk Masoor ILL6821
ILL 4605
1
ILL 5782
0.925
1
ILL 6024
0.895
0.902
1
Pant L 406
0.947
0.956
0.904
1
ILL 8117
0.926
0.892
0.883
0.935
1
ILL 7556
0.915
0.882
0.872
0.925
0.967
1
ILL 7715
0.895
0.882
0.872
0.904
0.924
0.934
1
ILL 6468
0.926
0.913
0.924
0.935
0.935
0.924
0.924
1
ILL 2580
0.905
0.892
0.903
0.935
0.935
0.924
0.903
0.914
1
Masoor 93
0.926
0.934
0.924
0.957
0.956
0.945
0.945
0.978
0.935
1
Turk Masoor
0.926
0.913
0.903
0.935
0.956
0.945
0.945
0.956
0.935
0.978
1
ILL6821
0.947
0.935
0.925
0.957
0.957
0.946
0.946
0.957
0.935
0.978
0.978
84
1
85
flowering (66 days), quick growing habit and has large seeds. Masoor 93 (Pakistan) takes 97 day to flower with spreading growth habit and plain/small seeds while Turk Masoor (collected from local market) takes 79 days to flower with spreading growth habit and spotted/medium large seeds. Red cotyledon colour is the only common character among these genotypes. Considering the contrasting phenologic and other morphological distinctness between Masoor 93 and Turk Masoor, crosses were made between these genotypes. In sub-cluster b, there was only one genotype namely ILL 6468 (in subcluster b, yellow cotyledon colour with plain/medium large seeds form ICARDA, Syria). This genotype flowers in 99 days with spreading growth habit and was crossed with ILL 7556 (in sub-cluster c). In sub-cluster c, 2 genotypes were grouped (ILL 7556 and ILL 8117 from India) and 96.7% similarity matrix was observed between them. Both the genotypes had spreading growth habit and red cotyledon colour but with spotted/small seeds (ILL 7556) and plain/small seeds (ILL 8117). In cluster C, three genotypes (ILL 4605 and ILL 5782 belonging to Argentina and ICARDA, Syria respectively, and Pant L 406 from India) were grouped forming two sub-groups (d and e). The genotype ILL 4605 (in sub-cluster e) is early flowering (81 days), has erect and quick seedling growth habit with yellow cotyledon colour, plain/large seeds and it was crossed with Pant L 406 (spreading growth habit, red cotyledon colour with spotted/ small seeds belonging to India).
Discussion The parental lines were evaluated for the assessment of genetic variation and to identify the major traits contributing to the variation for their potential use in the breeding programme. Significant variations in mean square values for different plant traits and a range of coefficients of variance (C. V.) implies variation in the key morphological characters among the parental lines. Therefore, hybridization among 86
combinations of these lines in the present study might be useful for the breeding programme to yield good results. Hybridization between any distantly related populations is expected to yield more heterotic and vigorous plants constituting much of the different traits contained in the two parental lines (Fikiru et al. 2007). Yield contributing characters such as plant height, productive branches, biomass and harvest index were observed more important traits to be used for the improvement of lentil seed yield. Phenology (days to flowering) and seed size of the parental lines was also observed to contribute significantly for the production of desirable segregants suitable to the environment. A number of studies are available (chapter 3) which emphasize on the consideration of yield related traits for the improvement in lentil seed yield as well as on the phenology of the plants and environment where the plants are grown. Most plant breeders assume that geographic patterns reflect genetic diversity but Jeena and Singh (2002), Kumar et al. (2004a) and (Yadav et al. 2005) could not find any direct associaition between geographic distribution and genetic divergence The studies of Ferguson et al. (1998a) did not reflect overall country relationships in the classification of macrosperma and microsperma types of lentil. The results of this study tend to support these findings whereas Lázaro et al. (2001) observed clear morphological groups based on seed characters. Seed size and other morphological characters had been found to form a continuum between macrosperma and microsperma types of lentil (Sohl and Erskine, 1984). The cluster analysis, generally, did not show clearly distinct geographic patterns except cluster B (ILL 7556 and ILL 2580, both belonging to India), and also irrespective grouping of the genotypes based on their seed size {(cluster A, ILL 8117 and Masoor 93, small-seeded while Turk Masoor, medium-seeded) and cluster D, ILL 7715, Pant L 406 and ILL 6468, small-seeded while ILL 5782 and ILL 6024, large-seeded)} might be attributed due to small size of the sample. The pattern of clustering obtained
87
seemed to be useful for selecting agronomically or morpho-metrically divergent parents in lentil crossing programme. In the present studies relating to the nature of inheritance in different plant/yield related traits, the combinations of parents used as male and female represent to different clusters validating the selection for crossing programme. In randomly amplified polymorphic DNA (RAPD) analysis, different levels of polymorphism were observed among the genotypes with several primers and none of the primers differentiated the genotypes. The higher levels of similarity observed among the genotypes might be an indicative of relatively low level of diversity in these genotypes but the genotypes had distinct phenological, growth habit, and other seed/cotyledon colour characteristics. Limited degree of genetic variability has been observed in cultivated lentils (Ahmad et al. 1996; Ford et al. 1997; Sharma et al. 1995; Sonnate and Pignone, 2001; Yüzbaşıoğlu et al. 2006). Similarly, morphological data, (Erskine et al. 1998), RFLPs (Havey and Muehlbauer, 1989) and the internal transcribed region (ITS) (Sonnate et al. 2003) indicated a lack of genetic diversity in Lens culinaris. Clustering of genotypes, in general, showed loose grouping based on geographic origin as well as on seed size (macrosperma and microsperma type) except for the grouping of ILL 2580 and ILL 7715 (both belonging to India having similar morphological and seed related characters showed the intervarietal genetic relationship relating to the country of origin). Clusters containing genotypes of heterogenous origin thereby indicating no parallelism of the clustering pattern in the grouping of lentil genotypes into close-knit clusters between genetic and geographic diversity have been observed in lentil (Chauhan et al. 2005; Poonam, 2006). Loose parallelism in the clustering pattern using RAPD markers has also been observed in A. paniculata (Lattoo et al. 2007). On the other hand, Iqbal et al. (1997) observed intervarietal genetic relationships of several cotton varieties related to the center of origin, and Sun et al. (2001) comparing microsatellite and RAPD
88
polymorphism in corn hybrids found that most of the hybrids from the same company were closely related to each other. The loose parallelism in clustering might be attributable due to random nature of the primers in this study which surveyed the genome and might involve the non-coding regions of the genome, and showed little conformity with the functional genome. There was no correspondence between genetic divergence and geographic origin as the genotypes from one origin dispersed randomly into more than one cluster. Tendency to yield such clustering pattern implies that the regional isolation may not always dilute the genetic make up of the introductions that contribute towards diversity in naturalized populations. Hence clustering seems to be influenced more by genetic constitution of the genotypes rather than the eco-geographic origin. This is possibly due to free exchange of seed material from the native places of wild abundance to other places for cultivation. The grouping of genotypes, used for the production of populations for inheritance studies of quantitative as well as qualitative traits, in different clusters based on morphological (Fig 4.1) as well as RAPD markers (Fig 4.3) shows good sampling of genotypes necessary for the crossing process as a source of variability. High similarity matrices among different genotypes showed relatively narrow genetic diversity in the parental lines. But morphologically, these genotypes were quite different for different plant/seed traits. It seemed that the primers did not relate according to their morphological characters in the genotypes. The irrespective grouping of macrosperma and microsperma types supported the findings of Abo-Elwafa et al. (1995) who observed that microsperma and macrosperma cultivars were indistinguishable by the RAPD markers. Despite the support of the above mentioned studies, the present RAPD analysis was informative enough for differentiation among the lentil parental lines.
89
Morphological evaluation as well as molecular marker analysis is recommended because they provide complementary information and increase the resolving power of genetic diversity analysis (Saha and Gopalakrishna, 2007). The evaluation of parents through morphological characters showed that yield related traits, and phenology suited to the environment of the plants may contribute significant role during selection and for formulating hybridization programmes. RAPD markers showed that relatively low genetic diversity existed among the genotypes. The clusters obtained through morphological characters as well as RAPD analysis, generally contained genotypes of heterogenous origin, thereby indicating no parallel between genetic and geographic patterns. Therefore, crosses between the members of clusters separated by inter-cluster distances are likely to yield desirable segregants of agronomic importance. These studies may provide valuable information to assist in parental selection for current and future lentil breeding programmes.
90
Chapter 5
Nature of Inheritance for Different Plant and Yield Related Traits Progress in varietal improvement in crop plants has been slow due to lack of imagination, vision and efficiency in utilizing, and to assess the components of genetic variation for various quantitative traits, which is likely to provide genetic basis for choosing parents for breeding programme (Krull and Borlaug, 1970). The choice of a suitable breeding programme depends upon the relative magnitude of different gene effects and an understanding of the mode of inheritance of complex quantitative characters. Emphasis has been put on the role of additive and dominance components in the inheritance of quantitative traits to formulate a breeding programme for improving a trait (Kunkaew et al. 2007). However, a number of researchers (Hayman, 1958; Brim and Cockerham, 1961; Stuber and Moll, 1974) have pointed the role of epistatic gene interactions in conditioning a character. Hill (1966) suggested the importance of epistatic interactions, even at digenic or trigenic level, in the inheritance of quantitative traits. Hence, the nature and magnitude of gene action involved in the expression of quantitative traits is important for successful development of crop varieties (Pradhan et al. 2006) and it is essential to know precisely the genetic architecture of character(s) for further improvements in seed yield. Among the various designs used, the generations mean analysis approach of Mather and Jinks (1982) is simple which provides the estimates of the main gene actions (additive and dominance) and their digenic {(i), (j), (l)} interactions, and helps in understanding the performance of the parents used in the
91
crosses and potential of the crosses to be used either for heterosis exploitation or pedigree selection. To collect information on the genetic behaviour controlling various plant traits in lentil (Lens culinaris Medik.), the generation means analysis was followed. The joint scaling tests for different plant characters were applied to test the adequacy of the additive-dominance model. The characters, where additive-dominance model was found inadequate (significant χ2 test), were subjected to further genetic analysis. Nonsignificant χ2 of joint scaling tests (Appendix-XXI and XXII) of the crosses PL 406 x ILL 4605 for days to flower, Turk Masoor x Masoor 93 for primary branches, ILL 5782 x ILL 2580 for nodes on primary and secondary branches, ILL 2580 x ILL 5782 for nodes on primary branches and ILL 7715 x ILL 6821 for seeds per pod indicated the adequacy of the additive-dominance model while the other crosses showing significant χ2 tests were subjected to further analysis for the detection of epistasis. Combined analysis of variance (Table 5.1) for twelve crosses showed highly significant differences in generations among all the traits studied. Partitioned analysis among different generations showed that highly significant differences were observed among parents for all the characters. In Ps vs F 1 s, highly significant differences were noted for days to flower, primary and secondary branches, nodes on primary branch, seeds per pod, pods per plant, 100 seed weight, biomass, harvest index and seed yield per plant while significant differences were observed in fertile nodes on primary branch, nodes on secondary branch and pod breadth. Among BC 1 s vs BC 2 s, days to flower differed highly significantly while days to mature showed significant differences. Highly significant differences were observed for days to flower, primary and secondary
92
Table 5.1: Mean square values for different plant characteristics in 12 crosses of lentil (Lens culinaris Medik.) Plant Primary Secondary Node on Fertile Node on Fertile Pod Pod Seed height Branch primary node on secondary node on length breadth pod-1 branch branch primary branch secondary plant-1 Plant-1 branch branch 0.683 0.0002 0.102 0.156 0.049 0.097 0.182 0.0002 0.0001 0.001
Source of variation d.f
Days to flower
Days To mature
Replication
2
0.065
1.398
Generation
5 58.909**
5.783** 16.090** 0.058** 13.963**
4.125** 3.997**
5.121**
3.912**
0.001** 0.002** 0.071** 3966.19** 0.197** 3.463** 24.303** 27.718**
P1s vs P2s
1 32.165** 22.554** 25.486** 0.027** 27.197**
4.329** 6.954**
8.124**
4.629**
0.002** 0.010** 0.275** 4976.70** 0.764** 4.672** 56.653** 17.559**
Ps vs F1s
1 183.847** 0.287
0.668
0.024** 11.953**
4.750** 0.253*
0.852*
0.114
0.0002
0.001*
0.039** 3137.98** 0.016** 0.469** 29.582** 20.167**
BC1s vs BC2s
1 16.570**
2.385*
0.033
0.0002
0.0009
0.001
0.014
0.271
0.015
0.0000
0.0001
0.0004
BCs vs F2s
1
0.002
1.195
0.130** 15.055**
0.331
2.327**
0.645*
4.851** 0.001** 0.001*
0.0001 5846.48** 0.0002
9.921** 0.001** 0.001*
0.039** 5865.42** 0.207** 8.725** 30.468** 59.541**
8.841**
100 Biomass Harvest plant-1 Index seed plant-1 weight
15.61
0.001
4.39
Ps, F1s vs BCs, F2s 1 53.121**
3.684** 53.070** 0.108** 15.611**
Error
10 0.328
0.253
0.245
0.002
0.103
0.118
0.039
0.097
0.038
0.0001
0.0001
0.001
11.63
Total
17 17.527
2.014
4.956
0.018
4.179
1.301
1.204
1.575
1.195
0.0003
0.0005
0.021
1175.20
** and *: Significant at P < 0.01 and P < 0.05 respectively P1s: All female parents, P2s: All male parents, BC1s: Backcrosses with female parents of all crosses, BCs: Both backcrosses (BC1s and BC2 s) of all crosses,
11.211** 10.439** 15.714**
Pods plant-1
Ps: Both female and male parents, F1s: F1generations of all crosses BC2s: Backcrosses with male parents of all crosses F2s: F2 generations of all crosses
93
0.0000
0.016
0.371
Seed yield plant-1 0.132
0.010
0.017
0.170
3.440**
4.795** 41.154**
0.0004 0.024
0.095
0.117
0.058
7.247
8.237
1.034
branches, fertile nodes on primary and secondary branch, pod length, pods per plant, biomass, harvest index and seed yield per plant in BCs vs F 2 s while nodes on secondary branch and pod breadth showed significant differences. Among uniform and segregating generations (Ps, F 1 s vs BCs, F 2 s), all the plant traits were noted highly significantly different except pod breadth where significant differences were observed. Character wise mean square values (analyses of variance) are presented in appendices IV-XX. Highly significant differences for days to flower were observed among generations in all the crosses (Appendix IV). Parents differed highly significantly in eight crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406, and PL 406 x ILL 4605) while significant differences were observed in two crosses (ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468). Highly significant differences were noted among Ps vs F1s in all the crosses except ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468. Among BC1s vs BC2s, highly significant differences were found in eight crosses (ILL 5782 x ILL 2580, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and PL 406 x ILL 4605). BCs vs F2s showed highly significant differences among five crosses (ILL 2580 x ILL 5782, ILL 7556 x ILL 6468, ILL 7715 x ILL 6821, ILL 4605 x PL 406, and PL 406 x ILL 4605). All the crosses, except ILL 6468 x ILL 7556, showed highly significant differences in Ps, F1s vs BCs, F2s. The estimates of additive, dominance and non-allelic interaction parameters for days to flower are presented in table 5.2. The estimates of mean (m) in all the crosses were significant. Both additive (d) and dominance (h) gene effects were significant in
94
Table 5.2: Estimates of additive, dominance and non-allelic gene effects for days to flower in 12 lentil crosses Cross
χ2
m
d
H
i
j
l
ILL 5782 x ILL 2580
102.05*±0.30
–2.97*±1.00
6.07*±2.38
-
–2.28*±1.02
–13.30*±3.57
32.71*
Duplicate
ILL 2580 x ILL 5782
99.63*±0.41
–1.15 ±1.05
17.97*±2.71
14.18*±2.66
-
–27.53*±3.84
36.09*
Duplicate
ILL 6024 x ILL 8117
100.81*±0.18
3.84*±0.66
9.35*±1.55
4.87*±1.51
3.48*±0.69
–12.90*±3.31
48.10*
Duplicate
ILL 8117 x ILL 6024
100.34*±0.25
4.48*±0.66
10.99*±1.69
6.49*±1.65
4.12*±0.68
–14.59*±3.36
48.63*
Duplicate
ILL 6468 x ILL 7556
98.75*±0.21
–5.00*±0.36
6.70*±1.16
-
-
–8.87*±2.99
115.11*
Duplicate
ILL 7556 x ILL 6468
97.55*±0.46
–4.92*±0.43
6.44*±2.05
6.42*±2.01
–4.27*±0.51
-
102.27*
-
ILL 6821 x ILL 7715
90.69*±0.74
–10.47*±2.59
25.99*±5.96
-
-
–27.70*±7.81
29.63*
Duplicate
ILL 7715 x ILL 6821
87.71*±1.01
–11.45*±2.62
38.58*±6.63
21.39*±6.62
-
–41.66*±8.08
23.37*
Duplicate
Masoor 93 x Turk Masoor
96.89*±0.26
–0.87 ± 0.49
3.99*±1.51
–3.60*±1.44
–9.82*±0.56
–12.90*±3.15
458.64*
Duplicate
Turk Masoor x Masoor 93
96.30*±0.32
–1.18*±0.46
6.92*±1.62
-
–10.13*±0.53
–17.10*±3.17
504.43*
Duplicate
ILL 4605 x PL 406
97.74*±0.79
–2.58 ±3.00
46.36*±6.84
21.89*±6.80
-
–30.10*±8.65
18.89*
Duplicate
PL 406 x ILL 4605
97.55*±0.02
–7.20*±0.30
41.33*±0.68
17.13*±0.61
–15.20*±0.31
–20.20*±2.87
291.33*
Duplicate
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
95
Type of interaction
nine crosses (ILL 5782 x ILL 2580, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Turk Masoor x Masoor 93 and PL 406 x ILL 4605). Additive gene effects were observed significantly positive in two crosses (ILL 6024 x ILL 8117 and ILL 8117 x ILL 6024) while in the seven crosses (ILL 5782 x ILL 2580, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Turk Masoor x Masoor 93 and PL 406 x ILL 4605), significantly negative effects were noted. Dominance (h) gene effect was significantly positive in all the crosses and had greater magnitude than that of additive gene effect (d). In three crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024 and PL 406 x ILL 4605), all the genetic parameters (additive, dominance and epistasis interactions) were significant. Dominance gene effect (h) had greater value than additive effect (d) in all the crosses. In three crosses (ILL 2580 x ILL 5782, Masoor 93 x Turk Masoor and ILL 4605 x PL 406), dominance gene effects were more pronounced. Among the digenic interaction effects, additive x additive (i) interactions were significant and positive in seven crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, ILL 7715 x ILL 6821, ILL 4605 x PL 406 and PL 406 x ILL 4605) while in cross Masoor 93 x Turk Masoor, significantly negative effect was observed. Positive and significant additive x dominance (j) gene effects were noted in two crosses (ILL 6024 x ILL 8117 and ILL 8117 x ILL 6024) while five crosses (ILL 5782 x ILL 2580, ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93 and PL 406 x ILL 4605) showed significantly negative effect. Significantly negative dominance x dominance gene effects (l) were noted in eleven crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) and
96
duplicate type of epistasis in these crosses was observed. In cross ILL 7556 x ILL 6468, the involvement of additive and non-additive gene interactions was observed. For days to mature (Appendix V), highly significant differences were observed in all the crosses for generations, P1s vs P2s (except for ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468) and Ps vs F1s. Highly significant differences were noted for BC1s vs BC2s in four crosses (ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, ILL 4605 x PL 406 and PL 406 x ILL 4605). Significant (ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468) and highly significant differences in eight crosses were found except two crosses (ILL 4605 x PL 406 and PL 406 x ILL 4605) for BCs vs F2s. In Ps, F1s vs BCs, F2s, highly significant differences in six crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 7556 x ILL 6468, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93), significant differences in two crosses (ILL 6468 x ILL 7556 and ILL 8117 x ILL 6024) were observed. The estimates of interacting gene effects for day to mature are presented in table 5.3. The estimates of mean (m) in all the crosses were significant. Significantly positive additive gene effect (d) was observed in two crosses (Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93) while four crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821) showed significantly negative effects. The dominance (h) component was significantly positive in six crosses (ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93) while significantly negative effect was observed in six crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468). Both additive (d) and dominance (h) gene effects were observed significant in six crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL
97
Table 5.3: Estimates of additive, dominance and non-allelic gene effects for days to mature in 12 lentil crosses Cross
j
l
χ2
m
d
H
i
ILL 5782 x ILL 2580
155.45*±0.20
–0.75±0.42
–13.22*±1.21
–1.50*±1.17
–3.05*±0.48
11.76*±3.08
364.70*
Duplicate
ILL 2580 x ILL 5782
155.48*±0.20
–0.50±0.44
–13.20*±1.22
–15.12*±1.19
–2.80*±0.49
12.08*±3.08
364.79*
Duplicate
ILL 6024 x ILL 8117
146.74*±0.26
–0.03±0.33
–20.02*±1.28
–15.5* ±1.22
4.34*±0.36
22.15*±3.04
289.99*
Duplicate
ILL 8117 x ILL 6024
146.47*±0.50
0.27±0.33
–18.34*±2.14
–14.16*±2.11
4.63*±0.35
21.19*±3.27
235.22*
Duplicate
ILL 6468 x ILL 7556
138.21*±0.16
–2.45*±0.31
–10.97*±0.92
–5.14*±0.90
–1.66*±0.33
7.02*±2.95
61.04*
Duplicate
ILL 7556 x ILL 6468
138.15*±0.15
–1.18*±0.35
–11.92*±0.98
–6.71*±0.93
-
11.63*±2.96
63.30*
Duplicate
ILL 6821 x ILL 7715 ILL 7715 x ILL 6821
132.47*±0.63 131.95*±0.63
–4.33*±1.24 –5.43*±1.24
15.33*±3.55 16.27*±3.55
9.91*±3.54 10.68*±3.54
-
-
41.08* 51.66*
-
Masoor 93 x Turk Masoor
132.00*±0.32
16.00*±0.37
19.89*±1.50
29.93*±1.48
8.74*±0.39
–32.40*±3.09
960.44*
Duplicate
Turk Masoor x Masoor 93
133.17*±0.32
15.92*±0.34
15.02*±1.48
25.35*±1.45
8.65*±0.37
–28.52*±3.08
879.91*
Duplicate
ILL 4605 x PL 406
138.59*±0.83
–1.22±1.70
16.37*±4.76
-
8.23*±1.70
11.96*±5.72
110.53*
Complementary
PL 406 x ILL 4605
139.40*±0.81
–1.93±1.63
12.36*±4.62
-
7.52*±1.63
15.20*±5.60
106.33*
Complementary
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
98
Type of interaction
6821, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93). Dominance gene effects were more pronounced in four crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117 and ILL 8117 x ILL 6024). In three crosses (ILL 6468 x ILL 7556, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93), all the genetic parameters (additive, dominance and epistasis interactions) were significant. Among the digenic interaction effects, additive x additive (i) interactions were negative and significant in six crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468 whereas in the four crosses (ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93), this interaction were found positive and significant. Positive and significant additive x dominance (j) gene effects were observed in six crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) while three crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782 and ILL 6468 x ILL 7556) showed significantly negative effects. Dominance x dominance (l) gene effects in eight crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 4605 x PL 406 and PL 406 x ILL 4605) were positive and significant while two crosses (Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93) showed negatively significant effects. Both additive and non-additive gene interactions were involved in two crosses (ILL 6821 x ILL 7715, ILL 7715 x ILL 6821). Duplicate type of gene action was observed in eight crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468) while two crosses (ILL 4605 x PL 406 and PL 406 x ILL 4605) showed complementary gene interactions.
99
Generations and parents showed highly significant differences for plant height in all the crosses (Appendix VI). In Ps vs F1s, cross ILL 7556 x ILL 6468 showed significant differences while highly significant differences were noted in other crosses except two crosses (ILL 5782 x ILL 2580 and ILL 2580 x ILL 5782). Highly significant differences in three crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024 and ILL 6468 x ILL 7556) and significant differences were found in two crosses (ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821) for BC1s vs BC2s generations. For BCs vs F2s, two crosses showed highly significant differences (ILL 8117 x ILL 6024 and ILL 7715 x ILL 6821) and the cross between ILL 2580 and ILL 5782 showed significant differences. Highly significant differences were found in eight crosses (ILL 5782 x ILL 2580, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) for uniform (Ps, F1s) vs segregating (BCs, F2s) generations while the cross between ILL 2580 and ILL 5782 showed significant differences. The estimates of interacting gene effects for plant height are presented in table 5.4. The estimates of mean (m) in all the crosses were significant. Additive gene effect (d) was observed significantly positive in five crosses (ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 7715 x ILL 6821 and Masoor 93 x Turk Masoor) while in three crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117 and ILL 6821 x ILL 7715), significantly negative effects were observed. In three crosses (ILL 2580 x ILL 5782, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821), significantly positive dominance effects (h) were observed while two crosses (ILL 6024 x ILL 8117 and Turk Masoor x Masoor 93) showed significantly negative effects. Significant gene effects of both additive (d) and dominance (h) components were observed in four crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821). More
100
Table 5.4: Estimates of additive, dominance and non-allelic gene effects for plant height in 12 lentil crosses Cross
i
d
H
ILL 5782 x ILL 2580 ILL 2580 x ILL 5782
35.98*±0.40 32.63*±0.60
1.43 ± 0.75 –1.50*±0.72
3.95 ± 2.33 10.47*±2.10
9.76*± 2.03
3.06*±0.83 -
-
44.95* 175.15*
-
ILL 6024 x ILL 8117 ILL 8117 x ILL 6024
32.26*±0.38 29.68*±0.33
–5.73*±0.93 5.58*±0.88
–4.90*±2.46 2.02 ±2.29
7.57*± 2.22
2.04*±0.97 13.35*±0.92
-
18.13* 312.73*
-
ILL 6468 x ILL 7556 ILL 7556 x ILL 6468
26.00*±0.32 29.48*±2.15
2.68*±0.51 2.32*±0.61
–2.78± 1.71 –15.04± 8.69
ILL 6821 x ILL 7715 ILL 7715 x ILL 6821
30.17*±0.37 28.38*±0.40
–3.43*±1.29 3.30*±1.27
24.72*±3.03 29.20*±3.06
Masoor 93 x Turk Masoor Turk Masoor x Masoor 93
26.14*±0.38 29.10*±0.32
2.12*±1.05 –0.88 ± 0.95
0.09 ± 2.64 –10.53*±2.33
ILL 4605 x PL 406 PL 406 x ILL 4605
24.14*±0.30 24.86*±0.36
0.25 ± 1.08 0.72 ± 1.08
4.73 ± 2.50 2.69 ± 2.63
-
j
l
χ2
m
Type of interaction
2.83*±0.58 7.83*±0.67
7.84*±3.28 22.22*±6.88
111.10* 218.43*
-
9.99* ±2.97 15.53*± 3.00
-
–12.07*±4.37
76.39* 85.28*
Duplicate
7.40*± 2.59 -
–2.68*±0.99
7.66*±3.76 15.74*±3.65
159.73* 59.24*
Duplicate
2.98*±1.11 3.44*±1.11
30.64*±3.72 35.19*±3.83
488.17* 419.73*
-
-
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
101
pronounced additive effects (d) were observed in four crosses (ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468 and Masoor 93 x Turk Masoor). Dominance effects (h) were more pronounced in cross Turk Masoor x Masoor 93. Among the digenic interactions, additive x additive (i) interactions was positive and significant in five crosses (ILL 2580 x ILL 5782, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and Masoor 93 x Turk Masoor). Positive and significant additive x dominance (j) gene effects were observed in seven crosses (ILL 5782 x ILL 2580, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 4605 x PL 406 and PL 406 x ILL 4605) while one cross (Turk Masoor x Masoor 93) showed negatively significant effects. Dominance x dominance gene effect (l) was found positively significant in six crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) whereas cross ILL 7715 x ILL 6821 showed negatively significant effects. Additive x additive (i) type of gene action was more pronounced in two crosses (ILL 2580 x ILL 5782 and ILL 6821 x ILL 7715) while two crosses (ILL 5782 x ILL 2580 and ILL 8117 x ILL 6024) showed more pronounced additive x dominance (j) type of gene interaction. Dominance x dominance (l) interaction was more prominent in six crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605). Duplicate type of epistasis was observed in two crosses (ILL 7715 x ILL6821 and Turk Masoor x Masoor 93) while other crosses showed additive and non-additive gene interactions. Primary branches (Appendix VII) showed highly significant differences for generations in all the crosses except cross ILL 7556 x ILL 6468 where significant differences were observed. One cross (Turk Masoor x Masoor 93) showed nonsignificant differences. For P 1 s vs P 2 s, six crosses (ILL 5782 x ILL
102
2580, ILL 2580 x ILL 5782, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821) showed highly significant differences. Ps vs F 1 s showed highly significant differences in seven crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93) and significant differences were noted in ILL 6468 x ILL 7556. In backcrosses (BC 1 s vs BC 2 s), highly significant differences were observed in two crosses (ILL 6024 x ILL 8117 and ILL 8117 x ILL 6024) while significant differences were found in cross ILL 6821 x ILL 7715. Highly significant differences were observed in seven crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7715 x ILL 6821 and Masoor 93 x Turk Masoor) while significant differences were found in cross between ILL 6821 and ILL 7715 for BCs vs F 2 s. Four crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 4605 x PL 406 and PL 406 x ILL 4605) in Ps, F 1 s vs BCs, F 2 s populations showed highly significant differences while significant differences were observed in three crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782 and Masoor 93 x Turk Masoor). The estimates of interacting gene effects for primary branches are presented in table 5.5. Non-significant estimates of χ 2 test in cross Turk Masoor x Masoor 93 showed that additive–dominance model was adequate to account for the total genetic variation in this trait and dominance gene effect (h) was more pronounced. The estimates of mean (m) in all the crosses were significant. Two crosses (ILL 8117 x ILL 6024 and ILL 7715 x ILL 6821) showed significantly positive additive gene effects (d) while in
103
Table 5.5: Estimates of additive, dominance and non-allelic gene effects for number of primary branches per plant in 12 lentil crosses Cross
m
D
h
i
j
l
χ2
Type of interaction
ILL 5782 x ILL 2580 ILL 2580 x ILL 5782
2.67*±0.03 2.55*±0.03
0.15±0.11 –0.08±0.12
1.115*±0.28 1.10*±0.30
0.90*±0.26 0.91*±0.28
0.37*±0.12 -
-
36.13* 47.17*
-
ILL 6024 x ILL 8117 ILL 8117 x ILL 6024
2.38*±0.03 2.33*±0.03
–1.02*±0.12 0.93*±0.12
1.34*±0.28 1.14*±0.28
1.44*±0.27 1.28*±0.27
0.97*±0.13 0.99*±0.13
-
87.99* 109.76*
-
ILL 6468 x ILL 7556 ILL 7556 x ILL 6468
2.29*±0.04 2.44*±0.03
–0.03±0.09 –0.02±0.11
1.40*±0.26 0.57 ±0.28
1.17*±0.23 -
–0.22*±0.11 -
-
57.66* 21.33*
-
ILL 6821 x ILL 7715 ILL 7715 x ILL 6821
2.10*±0.05 2.18*±0.05
–0.45*±0.13 0.35*±0.13
2.04*±0.34 2.88*±0.35
1.57*±0.33 2.31*±0.33
–0.15*±0.14 0.65*±0.14
-
46.12* 88.35*
-
Masoor 93 x Turk Masoor Turk Masoor x Masoor 93
2.34*±0.03 2.72*±0.04
0.07±0.12 0.02±0.04
1.11*±0.28 –0.19*±0.08
1.30*±0.27 -
-
-
40.29* 1.17
-
ILL 4605 x PL 406 PL 406 x ILL 4605
2.12*±0.04 2.14*±0.04
0.03±0.12 0.08±0.15
0.50±0.31 0.31±0.34
0.45*±0.29 -
-
-
55.64* 75.92*
-
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
104
two crosses (ILL 6024 x ILL 8117 and ILL 6821 x ILL 7715), significantly negative effects were observed. In nine crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and Masoor 93 x Turk Masoor), significantly positive dominance gene effects (h) were observed while in Turk Masoor x Masoor 93 showed significantly negative effects. The dominance effect (h) was significantly greater in magnitude than the additive component (d) in all crosses and was more pronounced in six crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93). Significant gene effects of both additive (d) and dominance (h) components were observed in four crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821). Among the digenic interactions, additive x additive (i) interaction was significantly positive in nine crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and ILL 4605 x PL 406). Positive and significant additive x dominance (j) gene effects were observed in four crosses (ILL 5782 x ILL 2580, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024 and ILL 7715 x ILL 6821) while two crosses (ILL 6468 x ILL 7556 and ILL 6821 x ILL 7715) showed negatively significant effects. Additive x additive (i) type gene action was more prominent in four crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, Masoor 93 x Turk Masoor and ILL 4605 x PL 406) and dominance gene effects (h) were more pronounced in cross ILL 7556 x ILL 6468. Two crosses (ILL 7556 x ILL 6468 and PL 406 x ILL 4605) showed significant joint scaling test but none of the gene effect or non-allelic interaction seems to be significant indicating the explanation difficult and gene action may be studied in F 3 or in further
105
generations. The remaining crosses showed the involvement of both additive and non-additive gene interaction. Ten crosses for secondary branches showed highly significant differences while two crosses (ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468) showed significant differences (Appendix VIII) in generations. Highly significant differences among parents (P 1 s vs P 2 s) were noted in eight crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93) while two crosses (ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468) showed significant differences. For Ps vs F 1 s, six crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, ILL 4605 x PL 406 and PL 406 x ILL 4605) showed highly significant differences. Backcrosses (BC 1 s vs BC 2 s) showed highly significant differences in two crosses (ILL 6024 x ILL 8117 and ILL 8117 x ILL 6024) while significant differences were found in three crosses (ILL 6468 x ILL 7556, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93). For BCs vs F 2 s, eight crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93 and PL 406 x ILL 4605) showed highly significant differences while significant differences were observed in cross ILL 4605 x PL 406. Seven crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) showed highly significant differences while in two crosses (ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468), significant differences were observed for Ps, F 1 s vs BCs, F 2 s generations.
106
The estimates of interacting gene effects for secondary branches are presented in table 5.6. The estimates of mean (m) in all the crosses were significant. Additive gene effect (d) was observed significantly positive in four crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 7715 x ILL 6821 and Masoor 93 x Turk Masoor) while in two crosses (ILL 6468 x ILL 7556 and Turk Masoor x Masoor 93), significantly negative effects were noted. Dominance gene effect (h) was significantly positive in all the crosses but this effect was more pronounced in six crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, ILL 4605 x PL 406 and PL 406 x ILL 4605). Significant gene effects of both additive (d) and dominance (h) were observed in six crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93). The dominance (h) component was significantly greater in magnitude than the additive gene effect (h) in eleven crosses except for cross ILL 6024 x ILL 8117. All the genetic parameters were observed significant in the cross Masoor 93 x Turk Masoor. Among the digenic interactions, additive x additive (i) interactions were significantly positive in all the crosses. Positive and significant additive x dominance (j) gene effects were observed in seven crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor, ILL 4605 x PL 406 and PL 406 x ILL 4605) while two crosses (ILL 6024 x ILL 8117 and Turk Masoor x Masoor 93) showed significantly negative effects. Dominance x dominance gene effect was found positively significant in two crosses (ILL 4605 x PL 406 and PL 406 x ILL 4605) while negatively significant effects were observed in five crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and Masoor 93 x Turk Masoor). The magnitude of additive x additive (i) interaction was greater than both the additive x dominance (j) and dominance x
107
Table 5.6: Estimates of additive, dominance and non-allelic gene effects for number of secondary branches per plant in 12 lentil crosses Cross
m
D
h
i
j
l
χ2
Type of interaction
ILL 5782 x ILL 2580
14.33*±0.34
–0.18±0.71
16.12*±2.09
11.64*±1.96
4.35*±0.79
–10.8*± 3.61
69.56*
Duplicate
ILL 2580 x ILL 5782
10.14*±0.27
–0.70±0.58
32.80*±1.71
29.12*±1.60
3.83*±0.67
–30.62*±3.28
450.30*
Duplicate
ILL 6024 x ILL 8117 ILL 8117 x ILL 6024
12.33*±0.33 10.28*±0.25
5.75*±0.90 6.18*±0.75
4.70*±2.28 12.36*±1.86
6.58*±2.24 13.58*±1.81
–2.42*±0.95 9.52*±0.81
-
58.36* 365.57*
-
ILL 6468 x ILL 7556 ILL 7556 x ILL 6468
13.19*±0.38 13.10*±0.31
–2.47*±0.76 1.70*±0.76
5.22*±2.22 5.63*±2.06
4.89*±2.15 4.93*±1.98
2.81*±0.81
-
22.19* 42.34*
-
ILL 6821 x ILL 7715
9.29*±0.30
–1.08±1.05
17.38*±2.46
11.14*±2.43
-
–9.66*±3.96
73.08*
Duplicate
ILL 7715 x ILL 6821
8.69*±0.25
–0.23±0.69
16.08*±1.74
11.11*±1.71
-
–9.72*±3.23
103.61*
Duplicate
Masoor 93 x Turk Masoor
9.36*±0.34
3.51*±0.63
14.83*±1.87
14.47*±1.84
1.71*±0.67
–8.44*± 3.28
194.86*
Duplicate
Turk Masoor x Masoor 93
11.06*±0.32
–2.05*±0.59
9.56*±1.76
9.39*±1.74
–3.85*±0.63
112.82*
-
ILL 4605 x PL 406
7.86*±0.28
1.15±0.91
13.15*±2.23
7.19*±2.14
6.21*±0.97
9.86*±3.71
239.13*
Complementary
PL 406 x ILL 4605
7.86*±0.24
0.01±0.88
13.62*±2.09
8.08*±2.00
5.06*±0.94
7.24*±3.54
239.47*
Complementary
-
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
108
dominance (l) interactions except for cross ILL 4605 x PL 406. In two crosses (ILL 4605 x PL 406 and PL 406 x ILL 4605), complimentary type of gene interactions were important while five crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and Masoor 93 x Turk Masoor) showed duplicate type of non-allelic interactions. The remaining crosses showed additive and non-additive gene interactions. Nodes on primary branch (Appendix IX) showed highly significant differences in eight crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93) while significant differences were observed in three crosses (ILL 2580 x ILL 5782, ILL 4605 x PL 406 and PL 406 x ILL 4605) among generations whereas for parents (P 1 s vs P 2 s), eight crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93) showed highly significant differences and four crosses
(Masoor 93 x Turk Masoor, Turk
Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) showed significant differences. In six crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93), highly significant differences were found whereas cross ILL 2580 x ILL 5782 showed significant differences for Ps vs F 1 s. BC 1 s vs BC 2 s showed significant differences in one cross ILL 6468 x ILL 7556 only. In BCs vs F 2 s, one cross (Turk Masoor x Masoor 93) showed highly significant differences while two crosses (ILL 5782 x ILL 2580 and ILL 7556 x ILL 6468) showed significant differences. Six crosses (ILL 6468 x ILL 7556,
109
ILL 7556 x ILL 6468, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93 and PL 406 x ILL 4605) showed highly significant differences while two crosses (ILL 8117 x ILL 6024 and ILL 4605 x PL 406) showed significant differences for Ps, F 1 s vs BCs, F 2 s generations. The estimates of interacting gene effects for nodes on primary branch are presented in table 5.7. Non-significant estimates of χ 2 test in crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782) showed that additive–dominance model was adequate to account for the total genetic variation in this trait while other crosses showed nonallelic interaction. In all the crosses, significant estimates of mean (m) were observed. In three crosses (ILL 5782 x ILL 2580, ILL 8117 x ILL 6024 and Masoor 93 x Turk Masoor), significantly positive while two crosses (ILL 6024 x ILL 8117 and ILL 6468 x ILL 7556) showed significantly negative additive gene effects (d). Significantly positive dominance gene effects (h) were observed in two crosses (ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821) while six crosses (ILL 2580 x ILL 5782, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93) showed significantly negative effects. Significant effects of both additive (d) and dominance (h) components were observed in three crosses (ILL 8117 x ILL 6024, ILL 6468 x ILL 7556 and Masoor 93 x Turk Masoor). All the genetic parameters were observed significant in cross ILL 8117 x ILL 6024 showing involvement of additive and non-additive gene effects. Among the digenic interactions, additive x additive (i) interactions were positive and significant in three crosses (ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and Masoor 93 x Turk Masoor) and negatively significant in four crosses (ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, Turk Masoor x Masoor 93 and ILL 4605 x PL 406). Positive and significant additive x dominance (j) gene effects were observed in four crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024,
110
Table 5.7: Estimates of additive, dominance and non-allelic gene effects for number of nodes on primary branch in 12 lentil crosses Cross
m
d
h
i
j
l
-
-
-
χ2
Type of interaction
ILL 5782 x ILL 2580 ILL 2580 x ILL 5782
21.94*±0.28 21.73*±0.26
0.57*±0.27 0.41±0.27
–0.72±0.52 –2.38*±0.50
ILL 6024 x ILL 8117 ILL 8117 x ILL 6024
20.13*±0.22 20.37*±0.23
–2.24*±0.61 1.97*±0.66
0.78 ±1.57 –5.28*±1.66
–4.41*±1.60
2.21*±0.67 6.41*±0.72
11.54*±3.27
30.74* 99.08*
Duplicate
ILL 6468 x ILL 7556 ILL 7556 x ILL 6468
17.71*±0.20 18.68*±0.20
–1.82*±0.54 0.87 ±0.64
–4.72*±1.42 –13.44*±1.56
–7.77*±1.51
2.82*±0.67
15.57*±3.30
12.69* 66.55*
Duplicate
ILL 6821 x ILL 7715 ILL 7715 x ILL 6821
18.94*±0.28 17.82*±0.18
–0.18±0.71 0.62±0.73
10.29*±1.88 7.20*±1.70
6.94*±1.81 3.23*±1.63
–2.38*±0.75 –1.58*±0.76
33.75* 93.94*
-
Masoor 93 x Turk Masoor
17.28*±0.22
1.35*±0.57
–11.96*±1.47
6.29*±1.44
18.54*±3.17
89.25*
Duplicate
Turk Masoor x Masoor 93
18.55*±0.20
–0.78 ±0.57
–15.13*±1.42
–9.30*±1.39
–1.55*±0.61
19.13*±3.14
66.10*
Duplicate
ILL 4605 x PL 406 PL 406 x ILL 4605
18.05*±0.21 16.00*±0.21
0.77±0.62 0.85±0.66
–2.97±1.58 2.47±1.65
–3.60*±1.51 -
1.78*±0.66 -
10.33*±3.23 -
26.20* 55.08*
-
-
-
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
111
2.48 5.62
-
ILL 7556 x ILL 6468 and ILL 4605 x PL 406) while in three crosses (ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and Turk Masoor x Masoor 93), significantly negative effects were observed. Dominance x dominance gene effect (l) was found significant in five crosses (ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, ILL Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93 and ILL 4605 x PL 406). The magnitude of dominance x dominance (l) interaction was greater than both the additive x additive (i) and additive x dominance (j) interactions in these crosses. The crosses where additivedominance model was adequate, additive gene effect (d) was more pronounced in cross ILL 5782 x ILL 2580 while dominance gene effect (h) was observed more prominent in cross ILL 2580 x ILL 5782). In cross ILL 6024 x ILL 8117, additive x dominance type of interaction (j) was observed more important. In cross ILL 6468 x ILL 7556, the joint scaling test was significant but none of the non-allelic component was significant and was difficult to explain. Likewise in cross PL 406 x ILL 4605, only the mean effect (m) was observed significant and gene action may be studied in the F 3 or further generations. Duplicate type of epistasis was noted in four crosses (ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93 and PL 406 x ILL 4605). Highly significant differences were observed in all the crosses for fertile nodes on primary branch (Appendix X) in generations as well as in parents (P 1 s vs P 2 s) except for four crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93) in P 1 s vs P 2 s. Ps vs F 1 s showed highly significant differences in nine crosses except for three crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782 and ILL 6024 x ILL 8117). Two crosses (ILL 6024 x ILL 8117 and ILL 8117 x ILL 6024) showed highly significant differences while two other crosses (ILL 7715 x ILL 6821 and
112
Masoor 93 x Turk Masoor) showed significant differences in backcrosses (BC 1 s vs BC 2 s). In segregating generations (BCs vs F 2 s), two crosses (ILL 5782 x ILL 2580 and ILL 2580 x ILL 5782) showed highly significant differences while in five crosses (ILL 8117 x ILL 6024, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and Masoor 93 x Turk Masoor), significant differences were noted. Ps, F 1 s vs BCs, F 2 s generations showed highly significant differences in seven crosses (ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) while in two crosses (ILL 6024 x ILL 8117 and ILL 7715 x ILL 6821), significant differences were found. The estimates of interacting gene effects for fertile nodes on primary branch are presented in table 5.8. The estimates of mean (m) in all the crosses were significant. Additive gene effects (d) were significantly positive in two crosses (ILL 8117 x ILL 6024 and Masoor 93 x Turk Masoor) while significantly negative effects were noted in four crosses (ILL 5782 x ILL 2580, ILL 6024 x ILL 8117, ILL 7556 x ILL 6468 and Turk Masoor x Masoor 93). Dominance gene effects in eight crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and PL 406 x ILL 4605) were significantly positive while the cross ILL 7556 x ILL 6468 showed strongly negative. Significant gene effects of both additive (d) and dominance (h) were observed in five crosses (ILL 5782 x ILL 2580, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 7556 x ILL 6468 and Masoor 93 x Turk Masoor). The dominance (h) component was significant and greater in magnitude than the additive effect (d) in nine crosses. Additive gene effect (d) was more important in cross Turk Masoor x Masoor 93 while dominance gene effect was more pronounced in four crosses (ILL 2580 x ILL 5782, ILL 6821 x ILL
113
Table 5.8: Estimates of additive, dominance and non-allelic gene effects for number of fertile nodes on primary branch in 12 lentil crosses Cross
χ2
m
d
h
i
j
l
ILL 5782 x ILL 2580
11.26*±0.21
–1.67*±0.57
12.54*±1.49
11.10*±1.42
-
–18.20*±3.22
60.86*
Duplicate
ILL 2580 x ILL 5782
9.61*±0.20
0.07 ±0.51
9.51*±1.36
9.63*±1.30
2.22*±0.57
–11.79*±3.10
86.48*
Duplicate
ILL 6024 x ILL 8117 ILL 8117 x ILL 6024
10.36*±0.22 9.47*±0.21
–3.55*±0.50 2.43*±0.46
3.56*±1.40 3.19*±1.29
4.86*±1.33 5.39*±1.25
5.00*±0.52
-
38.51* 166.60*
-
ILL 6468 x ILL 7556 ILL 7556 x ILL 6468
9.93*±0.22 10.66*±0.20
0.32 ±0.51 –0.98*±0.47
0.26 ±1.44 –9.09*±1.30
3.71*±1.35 –4.81*±1.24
-
12.94*±3.08
1081.83* 41.83*
Duplicate
ILL 6821 x ILL 7715 ILL 7715 x ILL 6821
10.41*±0.21 9.63*±0.20
–0.47±0.61 1.25±0.70
7.83*±1.55 7.57*±1.65
5.23*±1.48 4.86*±1.60
–1.36*±0.64 -
-
Masoor 93 x Turk Masoor Turk Masoor x Masoor 93
8.06*±0.21 8.75*±0.19
2.33*±0.42 –1.17*±0.45
4.01*±1.23 0.01 ±1.20
6.43*±1.20 -
1.67*±0.46 –1.83*±0.49
ILL 4605 x PL 406 PL 406 x ILL 4605
7.65*±0.19 6.80*±0.15
–0.02±0.67 –0.32±0.60
2.23 ±1.55 7.10*±1.37
4.70*±1.34
2.57*±0.69 2.27*±0.63
26.79* 53.81*
-
9.31*±3.01
2136.35* 240.62*
-
10.20*±3.21 -
148.77* 304.09*
-
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
114
Type of interaction
7715, ILL 7715 x ILL 6821 and PL 406 x ILL 4605). Among the digenic interactions, additive x additive (i) interactions were positive and significant in nine crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and PL 406 x ILL 4605) while in cross ILL 7556 x ILL 6468, significantly negative effects were observed. Positive and significant additive x dominance (j) gene effects were observed in five crosses (ILL 2580 x ILL 5782, ILL8117 x ILL 6024, Masoor 93 x Turk Masoor, ILL 4605 x PL 406 and PL 406 x ILL 4605) while two crosses (ILL 6821 x ILL 7715 and Turk Masoor x Masoor 93) showed negatively significant effects. Dominance x dominance gene effect (l) was found significantly positive in three crosses (ILL 7556 x ILL 6468, Turk Masoor x Masoor 93 and ILL 4605 x PL 406) while significantly negative effects were observed in two crosses (ILL 5782 x ILL 2580 and ILL 2580 x ILL 5782). Additive x additive type of gene action (i) was more pronounced in cross ILL 6468 x ILL 7556. Duplicate type of epistasis was observed in three crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782 and ILL 7556 x ILL 6468). Other crosses showed the involvement of additive and non-additive gene interactions. Nodes on secondary branch (Appendix XI) showed highly significant differences in generations. Parents (P 1 s vs P 2 s) showed highly significant differences in eight crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) and significant differences were observed in two crosses (ILL 6821 x ILL 7715, ILL 7715 x ILL 6821). Highly significant differences for Ps vs F 1 s generations were found in eight crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL
115
7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) while cross ILL 5782 x ILL 2580 showed significant differences. Backcrosses (BC 1 s vs BC 2 s) showed highly significant differences in four crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 7715 x ILL 6821 and Masoor 93 x Turk Masoor) and significant differences were noted in three crosses (ILL 6821 x ILL 7715, Turk Masoor x Masoor 93 and PL 406 x ILL 4605). BCs vs F 2 s showed highly significant differences in five crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821) and significant differences were noted in two crosses (ILL 6468 x ILL 7556 and Masoor 93 x Turk Masoor). Segregating generations (Ps, F 1 s vs BCs, F 2 s) showed highly significant differences in seven crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605). The estimates of interacting gene effects for nodes on secondary branch are presented in table 5.9. In cross ILL 5782 x ILL 2580, additive-dominance model was found adequate to account for the total genetic variation in this trait and significant additive (d) genetic effect showed greater importance. The estimates of mean (m) in all the crosses were significant. Additive gene effects (d) were significantly positive in six crosses (ILL 5782 x ILL 2580, ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and PL 406 x ILL 4605) while significantly negative effects were observed in three crosses (ILL 6024 x ILL 8117, ILL 6821 x ILL 7715 and Turk Masoor x Masoor 93). Dominance gene effects were significantly positive in five crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821) while three crosses (ILL 6468 x
116
Table 5.9: Estimates of additive, dominance and non-allelic gene effects for number of nodes on secondary branch in 12 lentil crosses Cross
m
d
h
i
j
l
χ2
Type of interaction
-
6.15 37.21*
-
–7.92*±3.13
61.09*
Duplicate
ILL 5782 x ILL 2580 ILL 2580 x ILL 5782
18.15*±0.25 16.83*±0.20
1.30*±0.25 0.20±0.61
–0.62±0.48 5.00*±1.56
5.80*±1.46
-
ILL 6024 x ILL 8117
15.75*±0.21
–3.10*±0.52
7.88*±1.39
7.96*±1.35
-
ILL 8117 x ILL 6024
15.26*±0.21
3.92*±0.60
8.49*±1.51
9.21*±1.47
8.10*±0.64
–9.74*±3.21
219.91*
Duplicate
ILL 6468 x ILL 7556
13.78*±0.18
–0.48 ±0.30
–12.32*±1.01
–7.70*±0.95
0.91*±0.35
23.45*±2.99
272.89*
Duplicate
ILL 7556 x ILL 6468
14.76*±0.18
1.42*±0.36
–16.73*±1.08
–12.34*±1.01
2.81*±0.40
29.29*±3.02
335.18*
Duplicate
ILL 6821 x ILL 7715
13.24*±0.27
–2.05*±0.82
15.04*±2.02
9.93*±1.97
–10.60*±3.50
49.54*
Duplicate
ILL 7715 x ILL 6821
11.73*±0.24
3.92*±0.70
17.98*±1.74
11.93*±1.69
4.98*±0.73
–6.68*±3.27
226.73*
Duplicate
Masoor 93 x Turk Masoor Turk Masoor x Masoor 93
12.68*±0.20 14.04*±0.19
2.55*±0.39 –1.62*±0.44
–0.69 ±1.14 –5.02*±1.20
4.98*±1.11 -
–4.12*±0.49
6.94*±3.05
214.84* 144.35*
Duplicate
ILL 4605 x PL 406 PL 406 x ILL 4605
13.03*±0.27 13.26*±0.25
–0.45 ±0.72 1.25*±0.62
–0.42±1.85 –1.02±1.68
–3.29*±1.61
1.87*±0.74 3.57*±0.65
12.85*±3.40 12.85*±3.26
56.83* 70.39*
-
-
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
117
ILL 7556, ILL 7556 x ILL 6468 and Turk Masoor x Masoor 93) showed significantly negative effects. More pronounced additive gene effects (d) were observed in three crosses (ILL 5782 x ILL 2580, Masoor 93 x Turk Masoor and PL 406 x ILL 4605) while two crosses (ILL 2580 x ILL 5782 and ILL 6468 x ILL 7556) showed that dominance gene effects (h) had more importance. Significant gene effects of both additive (d) and dominance (h) were observed in six crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and Turk Masoor x Masoor 93). All the genetic parameters were observed significant in three crosses (ILL 8117 x ILL 6024, ILL 7556 x ILL 6468 and ILL 7715 x ILL 6821). Among the digenic interactions, additive x additive (i) interactions were positive and significant in six crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and Masoor 93 x Turk Masoor) while three crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468 and PL 406 x ILL 4605) showed negatively significant effects. Positive and significant additive x dominance (j) gene effects were observed in six crosses (ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 7715 x ILL 6821, ILL 4605 x PL 406 and PL 406 x ILL 4605) whereas in cross Turk Masoor x Masoor 93, significantly negative effect was observed. Dominance x dominance gene effect (l) was found significantly positive in five crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) and the magnitude was greater than both of the additive x additive (i) and additive x dominance (j) interactions while negative effects were observed in four crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821). Additive x additive type of interaction (i) was more pronounced in crosses ILL 2580 x ILL 5782 and Masoor 93 x Turk Masoor. Dominance x dominance type of interaction was observed more pronounced in crosses ILL 4605 x
118
PL 406 and PL 406 x ILL 4605. Duplicate type of epistasis was observed in seven crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and Turk Masoor x Masoor 93). For fertile nodes on secondary branch (Appendix XII), generations showed highly significant differences in all the crosses. Parents (P 1 s vs P 2 s) showed highly significant differences in eight crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) while four crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821) showed significant differences. Eight crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) in Ps vs F 1 s showed highly significant differences while in one cross ILL 2580 x ILL 5782, significant differences were observed. In BC 1 s vs BC 2 s, three crosses (ILL 5782 x ILL 2580, ILL 6024 x ILL 8117 and ILL 8117 x ILL 6024) showed highly significant differences. Highly significant differences were found in seven crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and Masoor 93 x Turk Masoor) for BCs vs F 2 s and seven crosses (ILL 5782 x ILL 2580, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) in Ps, F 1 s vs BCs, F 2 s generations. Significant differences were noted in cross ILL 7715 x ILL 6821 in Ps, F 1 s vs BCs, F 2 s generations.
119
The estimates of interacting gene effects for fertile nodes on secondary branch are presented in table 5.10. The estimates of mean (m) in all the crosses were significant. Additive gene effects (d) were observed significantly positive in four crosses (ILL 2580 x ILL 5782, ILL 8117 x ILL 6024, ILL 7556 x ILL 6468 and ILL 7715 x ILL 6821) while three crosses (ILL 5782 x ILL2580, ILL 6024 x ILL 8117 and ILL 6468 x ILL 7556) showed significantly negative effects. Dominance gene effects were observed significantly positive in ten crosses while two crosses (ILL 6468 x ILL 7556 and ILL7556 x ILL 6468) showed significantly negative effects. Significant gene effects of both additive (d) and dominance (h) were observed in seven crosses (ILL 5782 x ILL2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468 and ILL 7715 x ILL 6821). Higher magnitude of dominance gene effects were observed in almost all the crosses except in crosses ILL 6468 x ILL7556 and ILL 7556 x ILL 6468 and were noted more pronounced in five crosses (ILL 6821 x ILL 7715, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605). All the genetic parameters were observed significant in five crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 8117 x ILL 6024, ILL 7556 x ILL 6468 and ILL 7715 x ILL 6821). Among the digenic interactions, additive x additive (i) interactions were positive and significant in eight crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93) while negatively significant in cross ILL 7556 x ILL 6468. Positive and significant additive x dominance (j) gene effects were observed in six crosses (ILL 2580 x ILL 5782, ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, ILL 7715 x ILL 6821, ILL 4605 x PL 406 and PL 406 x ILL 4605) whereas in two crosses (Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93), significant and negative effect was
120
Table 5.10: Estimates of additive, dominance and non-allelic gene effects for number of fertile nodes on secondary branch in 12 lentil crosses Cross
m
d
h
i
j
l
χ2
Type of interaction
ILL 5782 x ILL 2580
7.77*±0.19
–2.10*±0.41
9.09*±1.17
8.12*±1.13
–0.97*±0.45
–13.6* ±3.05
62.65*
Duplicate
ILL 2580 x ILL 5782
6.52*±0.18
0.78*±0.38
13.89*±1.09
12.42*±1.04
1.92*±0.43
–16.19*±3.01
175.25*
Duplicate
ILL 6024 x ILL 8117
6.97*±0.18
–3.17*±0.32
12.29*±1.05
12.18*±0.97
–15.10*±3.01
165.65*
Duplicate
ILL 8117 x ILL 6024
5.96*±0.16
3.03*±0.37
14.02*±1.00
14.77*±0.97
–18.00*±2.97
507.28*
Duplicate
ILL 6468 x ILL 7556
6.11*±0.17
–1.18*±0.29
–4.17*±0.98
-
7.22*±2.99
74.59*
Duplicate
ILL 7556 x ILL 6468
6.59*±0.16
0. 60*±0.26
–8.79*±0.90
–4.43*±0.84
1.43*±0.35
16.09*±2.96
180.26*
Duplicate
ILL 6821 x ILL 7715 ILL 7715 x ILL 6821
6.46*±0.19 5.30*±0.15
–0.87 ±0.50 1.05*±0.50
12.66*±1.30 16.27*±1.21
7.21*±1.26 11.37*±1.17
1.85*±0.53
–8.48*±3.06
108.88* 281.82*
Duplicate
Masoor 93 x Turk Masoor Turk Masoor x Masoor 93
5.08*±0.16 5.63*±0.12
–0.18±0.24 0.30±0.27
4.77*±0.82 2.11*±0.75
7.44*±0.80 4.76*±0.73
–2.11*±0.31 –1.63*±0.33
-
422.37* 410.45*
-
ILL 4605 x PL 406
4.48*±0.15
–0.18±0.49
2.74*±1.18
-
1.53*±0.51
11.20*±3.07
199.41*
Complementary
PL 406 x ILL 4605
4.96*±0.14
0.27±0.43
4.05*±1.07
-
1.98*±0.46
7.19*±3.03
140.16*
Complementary
5.75*±0.42 -
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
121
observed. Dominance x dominance gene effect (l) was found significantly positive in four crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 4605 x PL 406 and PL 406 x ILL 4605) while significantly negative in five crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024 and ILL 7715 x ILL 6821). Duplicate type of epistasis was observed in seven crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468 and ILL 7715 x ILL 6821) while complimentary type of gene interactions were observed in two crosses (ILL 4605 x PL 406 and PL 406 x ILL 4605). The involvement of additive and non-additive gene interactions in the remaining crosses was observed. Pod length showed highly significant differences of generations in ten crosses while in two crosses (ILL 5782 x ILL 2580 and ILL 2580 x ILL 5782), significant differences were noted (Appendix XIII). Among the parents, highly significant differences were found in ten crosses while two crosses (ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468) showed significant differences. Five crosses (ILL 6024 x ILL 8117, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 4605 x PL 406 and PL 406 x ILL 4605) were observed highly significantly different while five crosses (ILL 8117 x ILL 6024, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93) showed significant differences for Ps vs F 1 s. Highly significant differences were noted in two crosses (ILL 4605 x PL 406 and PL 406 x ILL 4605) for backcrosses (BC 1 s vs BC 2 s) while crosses ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821 showed significant differences. BCs vs F 2 s had highly significant differences in four crosses (ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468 and Turk Masoor x Masoor 93) and in two crosses (ILL 2580 x
122
ILL 5782 and ILL 7715 x ILL 6821), significant differences were found. Highly significant differences were found in four crosses (ILL 6468 x ILL 7556, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) while cross ILL 7715 x ILL 6821 showed significant differences for Ps, F 1 s vs BCs, F 2 s generations. The estimates of genetic parameters for pod length are presented in table 5.11. The estimates of mean (m) in all the crosses were significant. Additive gene effects (d) were significantly positive in two crosses (ILL 6821 x ILL 7715 and ILL 4605 x PL 406) while significantly negative effects were observed in crosses ILL 7715 x ILL 6821 and PL 406 x ILL 4605. Five crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468 and Turk Masoor x Masoor 93) showed significantly positive dominance gene effects (h) while in three crosses (ILL 2580 x ILL 5782, ILL 4605 x PL 406 and PL 406 x ILL 4605), significantly negative effects were observed. Significant gene effects of both additive (d) and dominance (h) were observed in three crosses (ILL 7715 x ILL 6821, ILL 4605 x PL 406 and PL 406 x ILL 4605). Additive gene effects were more pronounced in cross ILL 6821 x ILL 7556 while six crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468 and Turk Masoor x Masoor 93) showed more pronounced dominance gene effects (h). Among the digenic interactions, additive x additive (i) interactions were positive and significant in six crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7715 x ILL 6821 and Turk Masoor x Masoor 93) whereas in cross ILL 2580 x ILL 5782, significantly negative effects were found. Positive and significant additive x dominance (j) gene effects were observed in two crosses (Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93)
123
Table 5.11: Estimates of additive, dominance and non-allelic gene effects for pod length in 12 lentil crosses Cross
m
d
h
i
j
l
χ2
Type of interaction
ILL 5782 x ILL 2580 ILL 2580 x ILL 5782
1.03*±0.01 1.08*±0.01
–0.01±0.01 0.01±0.01
0.01 ±0.03 –0.20*±0.03
–0.20*±0.03
–0.05*±0.01 –0.03*±0.01
-
15.35* 63.29*
-
ILL 6024 x ILL 8117 ILL 8117 x ILL 6024
1.02*±0.01 1.01*±0.01
0.00±0.02 0.02±0.02
0.36*±0.05 0.40*±0.04
0.24*±0.04 0.33*±0.04
–0.08*±0.02 –0.06*±0.02
-
93.83* 87.38*
-
ILL 6468 x ILL 7556 ILL 7556 x ILL 6468
1.03*±0.01 1.01*±0.01
0.01±0.01 –0.01±0.01
0.31*±0.04 0.31*±0.04
0.28*±0.04 0.27*±0.04
-
62.35* 64.43*
-
ILL 6821 x ILL 7715 ILL 7715 x ILL 6821
1.15*±0.01 1.13*±0.01
0.04*±0.01 –0.05*±0.02
–0.05 ±0.05 0.17*±0.05
0.21*±0.04
–0.20*±0.02
-
8.53* 52.16*
-
Masoor 93 x Turk Masoor Turk Masoor x Masoor 93
1.10*±0.01 1.05*±0.01
–0.03±0.03 –0.01±0.02
0.03 ±0.07 0.14*±0.04
0.18*±0.04
0.17*±0.03 0.19*±0.02
-
64.00* 211.36*
-
ILL 4605 x PL 406 PL 406 x ILL 4605
1.05*±0.01 1.04*±0.01
0.07*±0.02 –0.07*±0.02
–0.18*±0.05 –0.13*±0.04
–0.08*±0.02 –0.28*±0.02
-
81.80* 421.80*
-
-
-
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
124
whereas in seven crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 7715 x ILL 6821, ILL 4605 x PL 406 and PL 406 x ILL 4605), significantly negative effect was observed. In three crosses (ILL 5782 x ILL 2580, Masoor 93 x Turk Masoor and ILL 4605 x PL 406), additive x dominance (j) interactions were more pronounced. In cross ILL 6821 x ILL 7715, it is difficult to explain gene action. Other crosses showed the involvement of additive and non-additive gene actions. Highly significant differences were observed in all the crosses for generations and parents for pod breadth (Appendix XIV). In eight crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and ILL 4605 x PL 406), highly significant differences were observed for Ps vs F 1 s. For backcrosses, six crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, ILL 4605 x PL 406 and PL 406 x ILL 4605) showed highly significant differences while significant differences were noted in ILL 8117 x ILL 6024, ILL 6468 x ILL 7556 and Masoor 93 x Turk Masoor crosses. BCs vs F 2 s showed highly significant differences in cross ILL 6024 x ILL 8117 while six crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, ILL 7715 x ILL 6821 and ILL 4605 x PL 406) showed highly significant differences. In six crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and PL 406 x ILL 4605), highly significant differences were found while four crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556 and PL 406 x ILL 4605) showed significant differences for Ps, F 1 s vs BCs, F 2 s.
125
The estimates of interacting gene effects for pod breadth are presented in table 5.12. The estimates of mean (m) in all the crosses were significant. Additive gene effects were significantly positive in four crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, Masoor 93 x Turk Masoor and ILL 4605 x PL 406) while two crosses (ILL 7715 x ILL 6821 and PL 406 x ILL 4605) showed significantly negative effects. Six crosses (ILL 5782 x ILL 2580, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, ILL 7715 x ILL 6821 and Turk Masoor x Masoor 93) showed significantly positive dominance gene effects while in cross ILL 4605 x PL 406, significantly negative effects were observed. Significant gene effects of both additive (d) and dominance (h) were observed in four crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 7715 x ILL 6821 and ILL 4605 x PL 406). Additive (d) gene effects were more pronounced in two crosses (Masoor 93 x Turk Masoor and PL 406 x ILL 4605) while dominance gene effects were more pronounced in three crosses (ILL 5782 x ILL 2580, ILL 7556 x ILL 6468 and Turk Masoor x Masoor 93). Among the digenic interactions, additive x additive (i) interactions were positive and significant in seven crosses (ILL 5782 x ILL 2580, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 7715 x ILL 6821 and Turk Masoor x Masoor 93). Positive and significant additive x dominance (j) gene effect was observed in cross Turk Masoor x Masoor 93 whereas in seven crosses (ILL 5782 x ILL 2580, ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor, ILL 4605 x PL 406 and PL 406 x ILL 4605), significantly negative effect was observed. Two crosses (ILL 2580 x ILL 5782 and ILL 6821 x ILL 7715) showed epistasis (significant χ2 test) but none of the non-allelic components was significant which is rather difficult to explain. In cross ILL 6468 x ILL 7556), additive x additive (i) type of gene interaction was more important while in two crosses (Masoor 93 x Turk Masoor and PL 406 x ILL 4605), additive x
126
Table 5.12: Estimates of additive, dominance and non-allelic gene effects for pod breadth in 12 lentil crosses Cross
m
d
h
i
j
l
χ2
Type of interaction
ILL 5782 x ILL 2580 ILL 2580 x ILL 5782
0.54*±0.01 0.57*±0.01
–0.01±0.01 –0.14±0.10
0.19*±0.03 0.30 ±0.21
0.14*±0.02 -
–0.03*±0.01 -
-
51.14* 48.19*
-
ILL 6024 x ILL 8117 ILL 8117 x ILL 6024
0.57*±0.01 0.58*±0.02
0.04*±0.01 0.02*±0.01
0.36*±0.02 0.25*±0.08
0.28*±0.02 0.18*±0.08
–0.08*±0.01
-
183.12* 87.38*
-
ILL 6468 x ILL 7556 ILL 7556 x ILL 6468
0.52*±0.01 0.51*±0.01
0.02±0.01 –0.02±0.01
0.03 ±0.02 0.10*±0.02
0.07*±0.02 0.10*±0.02
–0.03*±0.01
-
14.42* 95.74*
-
ILL 6821 x ILL 7715 ILL 7715 x ILL 6821
0.60*±0.01 0.58*±0.04
0.04 ±0.01 –0.07*±0.01
0.03 ±0.03 0.13*±0.03
0.10*±0.03
–0.10*±0.01
-
93.98* 127.43*
-
Masoor 93 x Turk Masoor Turk Masoor x Masoor 93
0.56*±0.01 0.54*±0.01
0.03*±0.01 –0.02 ±0.01
–0.02 ±0.02 0.07*±0.03
0.07*±0.03
–0.08*±0.01 0.03*±0.01
-
73.60* 21.51*
-
ILL 4605 x PL 406 PL 406 x ILL 4605
0.58*±0.01 0.56*±0.01
0.06*±0.01 –0.04*±0.01
–0.08*±0.03 0.02 ±0.03
–0.04*±0.01 –0.14*±0.01
-
9.94* 421.80*
-
-
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
127
dominance interaction (j) was more pronounced. Other crosses showed the involvement of additive and non-additive gene interactions. Generations, parents and Ps vs F1s showed highly significant differences in all the crosses for seeds per pod except three crosses for Ps vs F1s (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024 and ILL 6821 x ILL 7715) which showed nonsignificant differences (Appendix XV). Seven crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93 and ILL 4605 x PL 406) showed highly significant differences for backcross generations. For BCs vs F2s generations, five crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 8117 x ILL 6024, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93) showed highly significant differences while in cross ILL 4605 x PL 406, significant differences were observed. Uniform (Ps, F1s) vs segregating (BCs, F2s) generations showed highly significant differences in six crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) while in two crosses (ILL 6024 x ILL 8117 and ILL 6821 x ILL 7715), significant differences were observed. The estimates of interacting gene effects for seeds per pod are presented in table 5.13. Non-significant estimates of χ2 test in cross ILL 7715 x ILL 6821 showed that there was nothing beyond the additive–dominance model to account for the total genetic variation and dominance gene effect (h) was important in this trait having higher magnitude than additive gene effect (d). The estimates of mean (m) in all the crosses were significant. The additive gene effects (d) were observed significantly positive in five crosses (ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and PL 406 x ILL 4605) while significantly negative effects were observed in three crosses (ILL 6024 x ILL 8117, ILL 6468 x ILL 7556, Turk Masoor x
128
Table 5.13: Estimates of additive, dominance and non-allelic gene effects for seeds per pod in 12 lentil crosses Cross
m
d
h
i
j
l
χ2
Type of interaction
ILL 5782 x ILL 2580 ILL 2580 x ILL 5782
1.41*±0.02 150*± 0.01
–0.05±0.04 0.02±0.04
–0.89*±0.11 –1.13*±0.10
–0.54*±0.11 –0.81*±0.01
0.24*±0.05 0.31*±0.04
-
70.24* 122.48*
-
ILL 6024 x ILL 8117 ILL 8117 x ILL 6024
1.32*±0.04 1.21*±0.02
–0.21*±0.04 0.20*±0.04
0.20 ±0.19 0.59*±0.11
0.61*±0.10
–0.33*±0.05 -
-
43.98* 40.91*
-
ILL 6468 x ILL 7556 ILL 7556 x ILL 6468
1.57*±0.02 1.67*±0.02
–0.43*±0.05 0.39*±0.03
0.41*±0.13 –0.01 ±0.11
0.15*±0.05 0.68*±0.04
-
28.42* 334.73*
-
ILL 6821 x ILL 7715 ILL 7715 x ILL 6821
1.48*±0.02 1.65*±0.02
–0.09 ±0.05 0.11*±0.02
0.19*±0.13 –0.21*±0.03
0.28*±0.12 -
-
19.42* 7.57
-
Masoor 93 x Turk Masoor Turk Masoor x Masoor 93
1.40*±0.04 1.29*±0.02
0.11*±0.05 –0.16*±0.05
–0.45*±0.18 0.10 ±0.12
0.25*±0.12
0.51*±0.05 0.24*±0.05
-
96.88* 74.07*
-
ILL 4605 x PL 406 PL 406 x ILL 4605
1.18*±0.01 1.31*±0.02
–0.14*±0.06 0.11*±0.05
–0.02 ±0.13 –0.55*±0.13
0.33*±0.13 -
0.17*±0.06 0.42*±0.05
-
188.26* 180.47*
-
-
-
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
129
Masoor 93 and ILL 4605 x PL 406). Dominance gene effects (h) were significantly positive in four crosses (ILL 8117 x ILL 6024, ILL 6468 x ILL 7556 and ILL 6821 x ILL 7715) while significantly negative effects were observed in five crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and PL 406 x ILL 4605). The Significant gene effects of both additive (d) and dominance (h) were observed in five crosses (ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and PL 406 x ILL 4605). Additive gene effects (d) were more pronounced in four crosses (ILL 6024 x ILL 8117, ILL 7556 x ILL 6468, Turk Masoor x Masoor 93 and ILL 4605 x PL 406) while dominance effect (h) was more important in four crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6821 x ILL 7715 and PL 406 x ILL 4605). Among the digenic interactions, additive x additive (i) interactions were significantly positive in four crosses (ILL 8117 x ILL 6024, ILL 6821 x ILL 7715, Turk Masoor x Masoor 93 and ILL 4605 x PL 406) while negatively significant in two crosses (ILL 5782 x ILL 2580 and ILL 2580 x ILL 5782). Positive and significant additive x dominance (j) gene effect was observed in eight crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) whereas in cross ILL 6024 x ILL 8117, significant and negative effect was observed. Additive x additive (i) interaction was important in two crosses (ILL 8117 x ILL 6024 and ILL 6821 x ILL 7715). The additive x dominance (j) gene effect was more prominent in two crosses (ILL 6024 x ILL 8117, ILL 7556 x ILL 6468 and Turk Masoor x Masoor 93). In two crosses (Turk Masoor x Masoor 93 and ILL 4605 x PL 406), neither ‘h’ nor ‘l’ was significant and hence it is difficult to explain. Other crosses showed the involvement of additive and non-additive gene interactions.
130
Pods per plant (Appendix XVI) showed highly significant differences in four crosses (ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, ILL 4605 x PL 406 and PL 406 x ILL 4605) while in other eight crosses significant differences were noted in generations. Parents of half the crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, ILL 4605 x PL 406 and PL 406 x ILL 4605) showed highly significant differences while other half parents showed significant differences. Ps vs F 1 s showed highly significant differences in seven crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605). Three crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024 and ILL 7715 x ILL 6821) in backcrosses showed highly significant differences while two crosses (ILL 6821 x ILL 7715 and Turk Masoor x Masoor 93) had significant differences. In BCs vs F 2 s generations, eight crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and Turk Masoor x Masoor 93) showed highly significant differences while two crosses (ILL 5782 x ILL 2580 and Masoor 93 x Turk Masoor) showed significant differences. Cross ILL 6024 x ILL 8117 showed nonsignificant and two crosses (ILL 8117 x ILL 6024 and ILL 6468 x ILL 7556) showed significant differences while highly significant differences were observed in the remaining eight crosses for Ps, F 1 s vs BCs, F 2 s generations. The estimates of interacting gene effects for pods per plant are presented in table 5.14. The estimates of mean (m) were significant in all the crosses. Additive gene effects (d) were significantly positive in two crosses (ILL 8117 x ILL 6024 and ILL 7715 x ILL 6821) while two crosses (ILL 6024 x ILL 8117 and ILL 6821 x ILL 7715) showed
131
Table 5.14: Estimates of additive, dominance and non-allelic gene effects for pods per plant in 12 lentil crosses Cross
m
d
h
i
j
l
χ2
Type of interaction -
ILL 5782 x ILL 2580 ILL 2580 x ILL 5782
145.89*±4.76 99.56*±3.82
–16.70±10.60 –13.47± 9.15
116.35*±29.26 314.44*±24.53
321.95*±23.84
41.26*±11.15 44.49*± 9.78
–219.73*±24.06
129.79* 490.08*
Duplicate
ILL 6024 x ILL 8117
128.37*±5.12
–57.43*±11.50
303.88*±31.31
302.00*±30.81
-
–406.00*±23.70
111.69*
Duplicate
ILL 8117 x ILL 6024
82.57*±2.93
65.08*±11.05
530.94*±25.55
527.09*±25.03
107.47*±11.47
–668.75*±29.30
778.19*
Duplicate
ILL 6468 x ILL 7556
123.15*±4.68
13.32±9.56
297.47*±27.35
335.00*±26.76
41.04*±10.35
–404.00*±26.03
196.42*
Duplicate
ILL 7556 x ILL 6468
116.07*±3.90
–13.55±7.75
286.05*±22.80
323.89*±21.99
-
–354.24*±20.50
281.15*
Duplicate
ILL 6821 x ILL 7715 ILL 7715 x ILL 6821
73.90*±3.94 70.84*±3.27
–27.12*±12.63 40.80*±10.06
347.16*±30.30 372.12*±24.47
149.40*±29.78 162.72*±24.00
51.67*±10.16
-
303.33* 563.87*
-
Masoor 93 x Turk Masoor
90.29*±4.14
12.10±8.95
141.83*±24.89
131.20*±24.39
-
67.71*±26.59
309.38*
Complementary
Turk Masoor x Masoor 93
94.70*±3.43
–13.02±6.99
155.81*±20.09
141.75*±19.58
–34.03*±7.49
35.96*±17.93
400.76*
Complementary
ILL 4605 x PL 406
43.12*±2.25
–6.45±7.50
99.11*±18.12
39.75*±17.50
48.43*±7.76
227.80*±17.96
697.12*
Complementary
PL 406 x ILL 4605
46.85*±1.97
0.01±7.24
97.11*±16.99
33.32*±16.48
54.91*±7.51
234.63*±16.97
757.39*
Complementary
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
132
-
significantly negative effects. Dominance gene effects were significantly positive in all the crosses. The gene effects of both the additive (d) and dominance (h) were observed significant in four crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821). Dominance gene effects (h) were found more pronounced in eight crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605). The dominance (h) component was significantly greater in magnitude than the additive effect (d) in all the crosses. All the genetic parameters (additive, dominance, additive-additive, additive-dominance, and dominance-dominance) were observed significant in cross ILL 8117 x ILL 6024 indicating involvement of additive and non-additive gene effects. Among the digenic interactions, additive x additive (i) interactions were positive and significant in all crosses except for cross ILL 5782 x ILL 2580. Positive and significant additive x dominance (j) gene effects were observed in seven crosses (ILL 5782 x ILL 2580, ILL2580 x ILL 5782, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7715 x ILL 6821, ILL 4605 x PL 406 and PL 406 x ILL 4605) whereas in cross Turk Masoor x Masoor 93), significantly negative effect was observed. Dominance x dominance (l) gene effect was found positively significant in four crosses (Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) while negatively significant was observed in five crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468). Duplicate type of gene interaction was observed in five crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468) whereas complimentary type of gene interaction was observed in four crosses (Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605). The
133
remaining crosses showed the involvement of additive and non-additive gene interactions. 100 seed weight of ten crosses differed highly significantly in generations and in two crosses (ILL 6024 x ILL 8117 and ILL 8117 x ILL 6024) significant differences were noted (Appendix XVII). In parents, all crosses showed highly significant differences except for two crosses (ILL 6024 x ILL 8117 and ILL 8117 x ILL 6024) where nonsignificant differences were observed. Highly significant differences for Ps vs F 1 s generations were observed in eight crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 6821 x ILL 7715, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) while two crosses (ILL 8117 x ILL 6024 and ILL 7715 x ILL 6821) showed significant differences. Backcrosses (BC 1 s vs BC 2 s) showed highly significant differences in six crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6468 x ILL 7556, ILL 7715 x ILL 6821, ILL 4605 x PL 406 and PL 406 x ILL 4605) and significant differences were observed in cross ILL 6821 x ILL 7715. Five crosses (ILL 6468 x ILL 7556, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93) showed highly significant differences for BCs vs F 2 s while ILL 8117 x ILL 6024 showed significant differences. Ps, F 1 s vs BCs, F 2 s generations showed highly significant differences in all the crosses except for two crosses (ILL 6024 x ILL 8117 and ILL 8117 x ILL 6024) where nonsignificant differences were found. The estimates of interacting gene effects for 100 seed weight are presented in table 5.15. The estimates of mean (m) were significant in all the crosses. Significantly positive additive gene effects (d) were observed in four crosses (ILL 5782 x ILL 2580, ILL 6024 x ILL 8117, ILL 6821 x ILL 7715 and ILL 4605 x PL 406) while significantly
134
Table 5.15: Estimates of additive, dominance and non-allelic gene effects for 100 seed weight in 12 lentil crosses Cross
m
d
h
i
l
χ2
Type of interaction
0.19*±0.06 –1.61*±0.07
-
164.87* 1288.41*
-
j
ILL 5782 x ILL 2580 ILL 2580 x ILL 5782
2.59*±0.03 2.39*±0.03
0.90*±0.06 –0.89*±0.07
0.31 ±0.18 0.76*±0.18
ILL 6024 x ILL 8117 ILL 8117 x ILL 6024
2.79*±0.03 2.55*±0.03
0.14*±0.05 –0.11*±0.05
0.09 ±0.16 1.13*±0.17
0.85*±0.16
0.18*±0.05 -
-
14.83* 40.91*
-
ILL 6468 x ILL 7556 ILL 7556 x ILL 6468
2.02*±0.02 1.88*±0.02
0.05±0.04 –0.07±0.06
–0.22±0.12 0.27±0.14
–0.25*±0.11 0.29*±0.14
–0.22*±0.05 –0.35*±0.06
-
57.44* 169.30*
-
ILL 6821 x ILL 7715 ILL 7715 x ILL 6821
2.46*±0.03 2.32*±0.03
0.15*±0.05 –0.24*±0.04
1.43*±0.16 1.98*±0.14
1.25*±0.15 1.88*±0.14
–0.23*±0.05 –0.61*±0.05
-
105.16* 368.18*
-
Masoor 93 x Turk Masoor Turk Masoor x Masoor 93
2.41*±0.03 2.36*±0.02
0.01±0.04 –0.01±0.04
–2.04*±0.13 –1.90*±0.12
–1.52*±0.13 –1.39*±0.12
0.23*±0.04 0.22*±0.04
-
373.19* 408.62*
-
ILL 4605 x PL 406 PL 406 x ILL 4605
2.20*±0.03 2.31*±0.02
0.56*±0.11 –0.46*±0.11
–0.91 ±0.26 –1.44*±0.24
0.02*±0.26 –0.52*±0.24
–0.47*±0.12 –1.50*±0.11
-
51.68* 429.25*
-
-
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
135
negative effects were observed in four crosses (ILL 2580 x ILL 5782, ILL 8117 x ILL 6024, ILL 7715 x ILL 6821 and PL 406 x ILL 4605). Dominance gene effects (h) were significantly positive in four crosses (ILL 2580 x ILL 5782, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821) while three crosses (crosses Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93 and PL 406 x ILL 4605) showed significantly negative effects. Significant gene effects of both additive (d) and dominance (h) were observed in five crosses (ILL 2580 x ILL 5782, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and PL 406 x ILL 4605). In three crosses (ILL 5782 x ILL 2580, ILL 6024 x ILL 8117 and ILL 4605 x PL 406), additive component (d) was more pronounced while dominance gene effects were more pronounced in crosses Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93. Among the digenic interactions, additive x additive (i) interactions were positive and significant in five crosses (ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and ILL 4605 x PL 406) and in four crosses (ILL 6468 x ILL 7556, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93 and PL 406 x ILL 4605), significantly negative effects were observed. Positive and significant additive x dominance (j) gene effects were observed in four crosses (ILL 5782 x ILL 2580, ILL 6024 x ILL 8117, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93), whereas in seven crosses (ILL 2580 x ILL 5782, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, ILL 4605 x PL 406 and PL 406 x ILL 4605), significantly negative effect was observed. Additive x dominance (j) interactions were more pronounced in three crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782 and ILL 6024 x ILL 8117). Two crosses showed nonsignificant ‘h’ and ‘l’, indicating difficult to explain the gene interaction. The involvement of additive and non-additive gene interactions was observed in other crosses.
136
For biomass per plant, all the crosses showed highly significant differences in generations (Appendix XVIII). Parents showed highly significant differences in six crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 4605 x PL 406 and PL 406 x ILL 4605) and four crosses (ILL 6024 x ILL 8117 and ILL 8117 x ILL 6024, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821) showed significant differences. Ps vs F 1 s showed highly significant differences in nine crosses except for three crosses (ILL 6024 x ILL 8117, ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468) where nonsignificant differences were noted. In four crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 7715 x ILL 6821 and Turk Masoor x Masoor 93), highly significant differences were observed for backcrosses. BCs vs F 2 s showed highly significant differences in nine crosses except for three crosses (Masoor 93 x Turk Masoor, ILL 4605 x PL 406 and PL 406 x ILL 4605) where nonsignificant differences were observed. Highly significant differences were noted in seven crosses (ILL 2580 x ILL 5782, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) for Ps, F 1 s vs BCs, F 2 s and cross ILL 6821 x ILL 7715 showed significant differences. The estimates of interacting gene effects for biomass per plant are presented in table 5.16. The estimates of mean (m) in all the crosses were significant. Additive gene effects (d) were significantly positive in two crosses (ILL 5782 x ILL 2580 and ILL 8117 x ILL 6024) while three crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117 and Turk Masoor x Masoor 93) showed significantly negative effects. Dominance gene effects (h) in all the crosses were observed significantly positive except for cross Masoor 93 x Turk Masoor. The estimates of both the additive (d) and dominance (h) gene effects were
137
Table 5.16: Estimates of additive, dominance and non-allelic gene effects for biomass per plant in 12 lentil crosses Cross
m
d
h
i
j
l
χ2
Type of interaction
ILL 5782 x ILL 2580
12.37*±0.51
3.81*±0.96
9.65*±2.92
7.66*±2.80
5.92*±1.03
12.82*±4.01
168.84*
Complementary
ILL 2580 x ILL 5782
6.73*±0.27
–6.84*±0.77
29.15*±2.00
28.90*±1.88
–4.73*±0.86
–10.59*±3.49
745.59*
Duplicate
ILL 6024 x ILL 8117
8.86*±0.41
–5.17*±0.86
24.73*±2.44
23.54*±2.38
–4.14*±0.91
–32.30*±3.61
122.64*
Duplicate
ILL 8117 x ILL 6024
4.82*±0.20
4.22*±0.67
55.21*±1.67
51.18*±1.57
5.26*±0.74
–65.67*±3.25
1467.85*
Duplicate
ILL 6468 x ILL 7556
7.71*±0.35
1.04±0.80
26.57*±2.22
26.73*±2.13
2.78*±0.89
–21.70*±3.61
296.33*
Duplicate
ILL 7556 x ILL 6468
6.99*±0.26
0.09±0.51
26.53*±1.53
26.69*±1.46
1.67*±0.65
–18.77*±3.14
612.73*
Duplicate
ILL 6821 x ILL 7715
5.74*±0.30
–1.16±1.20
32.84*±2.75
18.35*±2.69
–8.52*±4.09
286.20*
Duplicate
ILL 7715 x ILL 6821
5.28*±0.29
1.72±0.98
37.10*±2.33
23.90*±2.27
2.44*±0.99
–20.41*±3.58
340.34*
-
Masoor 93 x Turk Masoor Turk Masoor x Masoor 93
6.50*±0.02 5.37*±0.24
1.07 ±0.63 –2.47*±0.61
–2.17 ±1.30 7.80*±1.62
9.19*±1.55
1.79*±0.68 –1.75*±0.65
18.29*±3.25 6.37*±3.19
338.49* 332.68*
Complementary
ILL 4605 x PL 406
3.11*±0.15
–0.24±0.37
4.73*±1.06
-
3.70*±0.41
22.81*±3.03
621.71*
Complementary
PL 406 x ILL 4605
3.52*±0.19
0.30±0.34
3.26*±1.11
-
4.24*±0.38
28.21*±3.05
781.73*
Complementary
-
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
138
significant in five crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024 and Turk Masoor x Masoor 93) and all other genetic parameters were also noted significant in these crosses. Dominance gene effect was more pronounced in six crosses (ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, ILL 4605 x PL 406 and PL 406 x ILL 4605). The dominance (h) component was significantly greater in magnitude than the additive effect (d) in all the crosses. Among the digenic interactions, significantly positive additive x additive (i) interaction was observed in nine crosses except for Masoor 93 x Turk Masoor, ILL 4605 x PL 406 and PL 406 x ILL 4605 crosses. Additive x dominance (j) gene effects were observed significantly positive in eight crosses (ILL 5782 x ILL 2580, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL7556 x ILL 6468, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor, ILL 4605 x PL 406 and PL 406 x ILL 4605) while three crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117 and Turk Masoor x Masoor 93) showed significantly negative effect. Dominance x dominance gene effect was found significantly positive in five crosses (ILL 5782 x ILL 2580, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) while seven crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821) showed significantly negative effects. Duplicate type of epistasis was observed in six crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468 and ILL 6821 x ILL 7715) while four crosses (ILL 5782 x ILL 2580, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) showed complimentary type of gene interactions. Additive and non-additive involvement of gene interactions was observed in two cross ILL 7715 x ILL 6821 and Masoor 93 x Turk Masoor.
139
Generations and parents showed highly significant differences in all the crosses for harvest index per plant (Appendix XIX). Significant (ILL 5782 x ILL 2580) and highly significant differences were observed in the crosses except for two crosses (ILL 2580 x ILL 5782 and ILL 6024 x ILL 8117) where nonsignificant differences were found for Ps vs F 1 s. In six crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821), highly significant differences were noted while cross ILL 8117 x ILL 6024 showed significant differences for backcrosses. BCs vs F 2 s generations showed highly significant differences in four crosses (ILL 2580 x ILL 5782, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93) and cross PL 406 x ILL 4605 showed significant differences. Eight crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) for Ps, F 1 s vs BCs, F 2 s showed highly significant differences. The estimates of interacting gene effects for harvest index per plant are presented in table 5.17. The estimates of mean (m) were significant in all the crosses. Additive gene effects (d) were significantly positive in three crosses (ILL 8117 x ILL 6024, ILL 7556 x ILL 6468 and ILL 7715 x ILL 6821) while two crosses (ILL 6468 x ILL 7556 and ILL 6821 x ILL 7715) showed significantly negative effects. Dominance gene effects were observed significantly negative in six crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605). Gene effects of additive (d) parameter were observed more effective in five crosses (ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821) whereas in six crosses (ILL 5782
140
Table 5.17: Estimates of additive, dominance and non-allelic gene effects for harvest index in 12 lentil crosses Cross
m
d
h
i
j
l
χ2
Type of interaction
ILL 5782 x ILL 2580
34.38*± 0.61
–0.72±0.92
–12.36*±3.08
–7.52*±3.03
-
7.96*±3.99
9.50*
Duplicate
ILL 2580 x ILL 5782
33.94*± 0.56
–0.27±0.95
–13.07*±2.97
–9.75*±2.93
-
17.23*±3.86
17.28*
Duplicate
ILL 6024 x ILL 8117
35.26*±0.47
–1.34 ±0.90
–4.27±2.66
-
-
9.58*±3.65
8.21*
-
ILL 8117 x ILL 6024
33.63*±0.55
2.22*±0.94
0.33±2.98
-
5.07*±1.00
30.32*
-
ILL 6468 x ILL 7556 ILL 7556 x ILL 6468
35.62*±0.66 34.41*±0.55
–5.23*±0.93 6.40*±1.14
–6.25±3.28 –2.14±3.24
-
–2.49*±1.01 9.14*±1.21
9.09*±4.26
8.09* 58.52*
-
ILL 6821 x ILL 7715 ILL 7715 x ILL 6821
30.78*±0.55 31.43*±0.54
–5.88*±1.40 4.03*±1.38
5.01±3.59 –5.38±3.54
–9.37*±3.50
–4.15*±1.46 5.76*±1.45
17.46*±4.45 34.37*±4.33
82.05* 104.08*
-
Masoor 93 x Turk Masoor
35.98*±0.82
0.35±1.14
–31.54*±4.08
–21.70*±4.00
3.95*±1.20
36.71*±4.62
58.10*
Duplicate
Turk Masoor x Masoor 93
33.69*±0.50
1.35±0.97
–26.89*±2.87
–15.86*±2.78
4.95*±1.04
31.84*±3.87
69.77*
Duplicate
ILL 4605 x PL 406
21.12*±0.43
–0.32±1.40
–8.46*±3.32
-
5.29*±1.44
22.33*±4.08
124.07*
Duplicate
PL 406 x ILL 4605
23.92*±0.82
–1.72±1.49
–15.78*±4.46
3.88*±1.53
25.30*±4.84
35.10*
Duplicate
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect,
–8.87*±4.43
-
h: Dominance gene effect, i: Additive x additive type of non-allelic interaction
j: Additive x dominance type of non-allelic interaction,
l: Dominance x dominance type of non-allelic interaction
141
x ILL 2580, ILL 2580 x ILL 5782, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605), dominance gene effect (h) was more pronounced. Among the digenic interactions, additive x additive (i) interactions were significantly negative in six crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93 and PL 406 x ILL 4605). Positive and significant additive x dominance (j) gene effects were observed in seven crosses (ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605) while two crosses (ILL 6468 x ILL 7556 and ILL 6821 x ILL 7715) showed significantly negative effect. Dominance x dominance gene effect (l) was found significant in ten crosses. In crosses ILL 8117 x ILL 6024 and ILL 6468 x ILL 7556, additive x dominance interaction (j) was more pronounced. Dominance x dominance (l) type of interaction was more pronounced in cross ILL 6024 x ILL 8117. Duplicate type of epistasis was observed in six crosses (ILL 5782 x ILL 2580, ILL 2580 x ILL 5782, Masoor 93 x Turk Masoor, Turk Masoor x Masoor 93, ILL 4605 x PL 406 and PL 406 x ILL 4605). Both additive and non-additive types of gene interactions were observed in other crosses. Seed yield per plant showed highly significant differences in all the crosses for generations as well as for parents except for two crosses (ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821) which showed significant differences for parents (Appendix XX). In five crosses (ILL 8117 x ILL 6024, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor and Turk Masoor x Masoor 93), highly significant differences were noted and cross ILL 7556 x ILL 6468 showed significant differences for Ps vs F 1 s. Highly significant differences were observed for backcrosses in six crosses (ILL 5782 x ILL 2580, ILL 2580 x
142
ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821) and two crosses (ILL 7556 x ILL 6468 and Turk Masoor x Masoor 93) showed significant differences. For BCs vs F 2 s, seven crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821) showed highly significant differences. Ps, F 1 s vs BCs, F 2 s generations showed highly significant differences in all the crosses except for two crosses (ILL 6024 x ILL 8117 and ILL 8117 x ILL 6024) which showed nonsignificant differences. The estimates of interacting gene effects for seed yield per plant are presented in table 5.18. The estimates of mean (m) in all the crosses were significant. Additive gene effects (d) were positively significant in four crosses (ILL 5782 x ILL 2580, ILL 8117 x ILL 6024, ILL 7556 x ILL 6468 and ILL 7715 x ILL) while significantly negative effects were observed in four crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 6821 x ILL 7715 and Turk Masoor x Masoor 93). Dominance gene effects were significantly positive in seven crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821) while the cross Masoor 93 x Turk Masoor
showed
significantly negative effects. Significant gene effects of both additive (d) and dominance (h) were observed in six crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715 and ILL 7715 x ILL 6821). More pronounced additive gene effects (d) were observed in two crosses (ILL 5782 x ILL 2580 and Turk Masoor x Masoor 93) while in two crosses (ILL 6468 x ILL 7556 and Masoor 93 x Turk Masoor), dominance gene effects (h) were more pronounced. All the genetic parameters were observed significant in three crosses (ILL 6024 x ILL 8117,
143
Table 5.18: Estimates of additive, dominance and non-allelic gene effects for seed yield per plant in 12 lentil crosses Cross
m
d
H
i
j
l
χ2
Type of interaction
202.43* 339.71*
-
ILL 5782 x ILL 2580 ILL 2580 x ILL 5782
4.18*±0.17 2.48*±0.13
1.34*±0.32 –2.11*±0.28
0.59 ±0.96 6.76*±0.78
7.57*±0.75
2.18*±0.42 –1.27*±0.39
ILL 6024 x ILL 8117
3.47*±0.02
–2.04*±0.34
6.82*±1.01
6.32*±0.99
–1.30*±0.36
–8.52*±2.96
52.01*
Duplicate
ILL 8117 x ILL 6024
1.67*±0.08
1.90*±0.27
18.72*±0.68
17.53*±0.64
2.63*±0.30
–22.34*±2.925
991.38*
Duplicate
ILL 6468 x ILL 7556
2.87*±0.15
–0.38 ±0.32
8.06*±0.89
8.71*±0.87
–7.37*±2.95
197.19*
Duplicate
ILL 7556 x ILL 6468
2.47*±0.10
0.86*±0.19
8.38*±0.59
9.24*±0.56
1.92*±0.23
–7.24*±2.88
473.75*
Duplicate
ILL 6821 x ILL 7715 ILL 7715 x ILL 6821
1.96*±0.12 1.85*±0.11
–0.95*±0.44 0.96*±0.37
11.25*±1.03 11.41*±0.90
5.60*±1.00 6.29*±0.87
1.28*±0.38
238.64* 290.27*
-
Masoor 93 x Turk Masoor
2.34*±0.11
0.34 ±0.23
–3.13*±0.64
–1.27*±0.63
1.09*±0.25
285.44*
Duplicate
Turk Masoor x Masoor 93
1.90*±0.10
–0.64*±0.19
–0.44 ±0.57
1.43*±0.55
333.19*
-
ILL 4605 x PL 406 PL 406 x ILL 4605
0.68*±0.04 0.77*±0.04
–0.05±0.12 0.05±0.10
0.42±0.31 –0.02±0.27
890.14* 1175.56*
-
-
-
1.52*±0.14 1.62*±0.11
6.54*±3.00 -
8.71*±2.90 6.57*±2.85 7.68*±2.84
*: Significant at P < 0.05 m: Mean effect, d: Additive gene effect, h: Dominance gene effect, i: Additive x additive type of non-allelic interaction j: Additive x dominance type of non-allelic interaction, l: Dominance x dominance type of non-allelic interaction
144
ILL 8117 x ILL 6024 and ILL 7556 x ILL 6468). Among the digenic interactions, additive x additive (i) interactions were positive and significant in eight crosses (ILL 2580 x ILL 5782, ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468, ILL 6821 x ILL 7715, ILL 7715 x ILL 6821 and Turk Masoor x Masoor 93) while significantly negative effects were observed in cross Masoor 93 x Turk Masoor. Positive and significant additive x dominance (j) gene effects were observed in seven crosses (ILL 5782 x ILL 2580, ILL 8117 x ILL 6024, ILL 7556 x ILL 6468, ILL 7715 x ILL 6821, Masoor 93 x Turk Masoor, ILL 4605 x PL 406 and PL 406 x ILL 4605) while two crosses (ILL 2580 x ILL 5782 and ILL 6024 x ILL 8117) showed significantly negative effects. Dominance x dominance gene effect was found significantly positive in four crosses (ILL 5782 x ILL 2580, Masoor 93 x Turk Masoor, ILL 4605 x PL 406 and PL 406 x ILL 4605) while significantly negative effects were observed in four crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556 and ILL 7556 x ILL 6468). Cross Turk Masoor x Masoor 93 showed more pronounced additive x additive (i) type of gene interaction. Three crosses (ILL 5782 x ILL 2580, ILL 4605 x PL 406 and PL 406 x ILL 4605) showed more effects of dominance x dominance (l) interaction. Opposite sign of (h) and (l) genetic parameters showed duplicate type of epistasis in five crosses (ILL 6024 x ILL 8117, ILL 8117 x ILL 6024, ILL 6468 x ILL 7556, ILL 7556 x ILL 6468 and Masoor 93 x Turk Masoor). Other crosses showed the involvement of additive and non-additive gene interactions. Summarizing the results, a simple additive-dominance model was adequate as inferred from the joint scaling test (nonsignificant χ2 test) for primary branches in Turk Masoor x Masoor 93, nodes on primary branch in two crosses (ILL 5782 x ILL 2580 and ILL 2580 x ILL 5782), nodes on secondary branch in ILL 5782 x ILL 2580, and seeds per pod in ILL 7715 x ILL 6821. Of these, dominance gene effect (h) was more
145
prominent in three crosses (Turk Masoor x Masoor 93 for primary branches, ILL 2580 x ILL 5782 for nodes on primary branch and ILL 7715 x ILL 6821 for seeds per pod) while more pronounced additive gene effect (d) were observed for nodes on primary and secondary branches in ILL 5782 x ILL 2580. For the remaining crosses and characters, digenic epistasis interactions were observed. Duplicate type of gene interactions were observed more prevalent in most of the crosses for different plant characters followed by complementary type of gene interactions. For days to flower and mature, duplicate type of gene interactions were observed more prominent. Plant height showed dominance x dominance interaction (l) more prominent followed by additive (d) and non-additive (h) type of gene interactions, and additive x dominance (j) interaction. Both additive and non-additive type of gene interactions seemed more influential in primary branches while secondary branches showed duplicate as well as additive and non-additive type of gene interactions. Nodes on primary branches showed duplicate type of gene interaction followed by additive and non-additive type of gene interactions while duplicate followed by dominance x dominance interactions (l) were observed more prevalent for nodes on secondary branch. Fertile nodes on primary and secondary branches showed more prominent effects of duplicate as well as additive and non-additive type of gene interactions. Additive and non-additive type of gene interactions along with additive x dominance (j) interactions were more important in pod length and breadth, seeds per pod, and 100 seed weight characters. Duplicate as well as complementary type of gene interactions were more pronounced for pods and biomass per plant. Harvest index was observed under the influence of duplicate type of interaction followed by additive x dominance (j) interaction. Duplicate as well as dominance x dominance (l) type of interactions were more prominent for seed yield per plant followed by additive and nonadditive type of gene interactions.
146
Discussion For effective selection, information on genetic parameters associated with inheritance of the character is prerequisite for planning a good and sound breeding programme. In lentil, both additive and non-additive gene effects were observed important in different studies for number of branches per plant (Sagar and Chandra, 1980; Sandhu et al. 1981; Waldia and Chhabra, 1989), time to flowering and maturity, and plant height (Swarup et al. 1991), seeds per pod (Jain, 1971) biological yield, plant spread and plant height (Chauhan and Singh, 1996), pods per plant (Waldia et al. 1989) and seed yield per plant (Singh and Jain, 1971; Waldia et al. 1989). Differences among generations as well as parents indicate the presence of genetic variability in the materials and diversity in the parents used. Kunkaew et al. (2007) observed significant differences among generations in azuki bean for seed yield per plant. In melon (Zalapa et al. 2006), significant differences were observed among generations and parents indicating the genetic variability in the materials used. Mean square values for all the plant traits in the present studies showed significant differences among generations as well as among the parents (Table 5.1) revealing sufficient genetic variability in the materials used. Gene effects for different plant characteristics among the crosses indicated that mean effect (m) of each cross was significant revealing differences in the traits among the parents which are in line to those observed in cotton (Esmail, 2007; Khan et al. 2007), winter wheat (Hakizimana et al. 2004), azuki bean (Kunkaew et al. 2007), cluster bean (Mitra et al. 2001), durum wheat (Sharma et al. 2003), and pea (Singh and Sharma, 2001). Akhtar and Chowdhry (2006) reported that most of the plant traits in bread wheat exhibited simple inheritance with additive-dominance model and several other researchers observed simple additive-dominance model sufficient to explain the genetics 147
of number of primary branches and plant height in chickpea (Kidambi et al. 1990), in sesame (Ganesh and Sakila, 1999; Sharmila et al. 2007) for plant height, capsule length, number of capsules and seed yield per plant, in Vigna sesquipedalis Frauw for seeds per pod and seed weight (Rahman and Saad, 2000) and in cotton for boll weight (Esmail, 2007). The crosses where χ2 tests were significant revealed that the epistatic interaction of genes had played a significant role in controlling the inheritance of these traits and the additive-dominance model was not sufficient to explain the genetic inheritance. In the present studies, a simple additive-dominance model was observed adequate in different crosses for different characters (primary branches in Turk Masoor x Masoor 93, nodes on primary branch in two crosses ILL 5782 x ILL 2580 and ILL 2580 x ILL 5782, nodes on secondary branch in ILL 5782 x ILL 2580, and seeds per pod in ILL 7715 x ILL 6821). Dominance gene effect (h) was more prominent for days to flower, primary branches, nodes on primary branch and seeds per pod in three crosses (Turk Masoor x Masoor 93, ILL 2580 x ILL 5782 and ILL 7715 x ILL 6821 respectively) while more pronounced additive gene effect (d) was observed for nodes on primary and secondary branches in cross ILL 5782 x ILL 2580 indicating the use of pedigree and heterosis breeding methods for the traits under the influence of ‘d’ and ‘h’ components respectively. Rahman and Saad (2000) in Vigna sesquipedalis Frauw and Singh and Singh (1990) in pea also suggested pedigree and heterosis breeding methods where‘d’ and ‘h’ components respectively were prominent. Predominant additive gene effects (d) have been observed for days to first flower, plant height, primary branches per plant and seeds per pod in lentil (Kumar et al. 1999). In Vigna sesquipedalis Frauw, Rahman and Saad (2000) reported that seeds per pod and seed weight were governed by additive gene effect, boll weight in cotton (Esmail, 2007) and number of capsules in sesame (Ganesh and Sakila, 1999) while dominance effect (h) in lentil was found to be more prominent
148
for secondary branches per plant, pods per plant, 100 seed weight and yield per plant (Kumar et al. 1999), seed yield in azuki bean (Kunkaew et al. 2007), seed yield in soybean (Kunta et al. 1997) and in peanut (Jogloy et al. 1999), and leaf breadth in cluster bean (Mitra et al. 2001). Ganesh and Sakila (1999) suggested biparental mating and heterosis breeding approaches in sesame where additive (d) and dominance (h) gene effects respectively were predominant. Esmail (2007) suggested pedigree selection method in cotton to improving the populations where lower magnitude of additive gene effects (d) relative to the corresponding dominance effects (h) were observed. However, the negative value of (h) observed for the traits in the respective crosses indicated that the alleles responsible for less value of the trait were dominant over the alleles controlling high value. For the traits in crosses where additive (d) gene effects were observed more pronounced, selection in early segregating generation may prove useful following biparental mating and for prominent dominance (h) gene effect, selection in late generation following heterosis technique may be effective. Greater estimates of dominance effect (h) than additive effect (d) indicate that the parents used in the present studies were diverse for the concerned traits and epistatic interaction of additive x additive (i), additive x dominance (j) and dominance x dominance (l) were integral components of the genetic formation of these traits. Hence, estimation and consideration of this gene effect component (epistasis) are important to formulating the lentil breeding programme. The importance of additive and non-additive gene effects for seed size and seed yield (Chauhan and Singh, 1995a), biological yield, plant spread and plant height in lentil (Chauhan and Singh, 1996), and for seed yield in azuki bean has been suggested (Kunkaew et al. 2007). Dominance gene effect (h) was predominantly prevalent and had higher magnitude for all the traits studied except for plant height, seeds per pod, 100 seed weight and harvest index per plant where
149
prevalence of both the additive (d) and dominance (h) gene effects in the respective crosses was observed more or less at par with each other indicating the importance of both the additive and dominance components in these crosses. The presence of nonadditive gene action is a limiting factor to exercise selection for evolving pure line. In such a situation, maximum gain could be achieved by maintaining considerable heterozygosity through mating of selected plants in segregating generation or by following some forms of recurrent selection (Compton, 1968; Parlevliet and van Ommeren, 1988). The additive, dominance and epistatic types of gene interactions in each cross for different traits were found to be different from each other and relative magnitude of these gene actions for the characters in each cross varied, leading to the variation in the inheritance. The dominance x dominance (l) interaction was larger than either additive x additive (i) or additive x dominance (j) effects and /or both for different traits, while for the main effects the dominance component (h) was greater than the additive (d) component in most of the crosses for different plant characters. The dominance (h) and dominance x dominance (l) effects were in the opposite direction, suggesting the prevalence of duplicate type epistasis in most cases which indicated predominantly dispersed alleles at the interacting loci (Jinks and Jones, 1958). Partitioning of total epistasis revealed that dominance x dominance (l) interaction was larger than either additive x additive (i) or additive x dominance (j) interactions in majority of the crosses for days to flower and mature indicating predominant duplicate type of gene interactions for these traits and heterosis breeding might be effective for the development of superior populations (Singh and Singh, 1990). Both additive (d) and dominance (h) components were observed more important for plant height. Among the digenic interactions, it showed dominance x dominance
150
interaction (l) more prominent in different crosses. Although additive (d) gene effects were significant for this trait, non-additive gene effects appeared to have been more important. Thus, hindrance may occur in selection improvement. Recurrent selection is likely to be useful for effective utilization of both the additive and non-additive gene effects prevalent in this character simultaneously. Kidambi et al. (1990) observed simple additive-dominance model sufficient to explain the genetics of plant height and number of primary branches in chickpea while digenic interactions were noticed for number of secondary branches, number of days to flowering and maturity. Additive and non-additive type of gene interactions seemed more influential in primary branches followed by additive x additive interaction (i) except for one cross where in a simple additive-dominance model, ‘h’ was prominent suggesting that selection should be postponed till the late generations through heterosis breeding. For other crosses, further generations are needed to study the gene action. For crosses where none of the non-allelic interaction was observed significant for the traits was difficult to explain. The same type of results were observed in bread wheat for grain yield per plant and suggested F3 or further generations to study the gene effect (Akhtar and Chowdhry, 2006 ). Secondary branches showed duplicate as well as additive and non-additive type of gene interactions followed by complementary gene effects. Nodes on primary branches showed duplicate type of gene interaction followed by additive and nonadditive type of gene interactions. Duplicate followed by dominance x dominance interaction (l) was observed more prevalent for nodes on secondary branch. Fertile nodes on primary and secondary branches showed more prominent effects of duplicate as well as additive and non-additive type of gene interactions. Additive and non-additive type of gene interactions along with additive x dominance (j) interactions were more important
151
in pod length and breadth, and seeds per pod characters. Duplicate as well as complementary type of gene interactions were more pronounced in pods per plant and biomass per plant. Harvest index was observed under the influence of duplicate type of interaction followed by additive x dominance (j) interaction. Kidambi et al. (1990) observed additive and non-additive type of gene interactions in chickpea for seed yield per plant and its components. Duplicate type of interaction shows complex nature of inheritance and may tend to hinder the progress making it difficult to fix the genes at increased level of manifestation. Jagtap (1986) and Abo El-Zahab (2000) in cotton suggested the possibility of obtaining desirable segregates through inter-mating in early generations by breaking undesirable linkage or to adopt recurrent selection for rapid improvement, when epistatic effects were more pronounced while Ramalingam and Sivasamy (2002) suggested delayed selection and inter-mating the segregants followed by recurrent selection for improvement of the trait when predominance of additive x dominance (j) epistatic effect (highest magnitude) for the trait were prevalent. For complementary type of gene action of certain traits in cluster bean, Mitra et al. (2001) suggested the possibility of considerable amount of heterosis in those crosses, and hybrid breeding and reciprocal recurrent selection methods were proposed for developing elite population. More pronounced effects of additive (d), additive x additive (i) and additive x dominance (j) type gene interactions indicated that seed weight trait was under the influence of both fixable and non-fixable gene effects suggesting the possibility of obtaining transgressive segregants in later generations and intermating the segregants followed by recurrent selection. Chauhan and Singh (1995b) suggested multiple factors with incomplete dominance responsible for seed size in lentil. They (1997) also observed preponderance of additive genetic variance for 100 seed weight.
152
Duplicate as well as dominance x dominance (l) type of interactions were more prominent for seed yield per plant followed by additive and non-additive type of gene interactions. Seed yield per plant was controlled by genes with significant additive, dominance, and epistatic effects suggesting that an effective selection to improve this trait should be mild in earlier generations and intensive in later generations. Chauhan and Singh (1997) observed non-additive genetic variance for seed yield in lentil. Dominance and epitasis gene action have been reported more prominent in rice by Shrivastava and Seshu (1983) and Dwivedi et al. (1999). Duplicate type of non-allelic interaction has been observed by Esmail (2007) in cotton for seed cotton and lint yield per plant, boll number and days to first flower. The same types of findings were also reported by Tandon et al. (1968) in barley. Ahuja and Dhayal (2007) observed preponderance of non-additive gene action in the inheritance of seed cotton yield and majority of its components. Gangappa et al. (1997) found predominant dominance control for seed yield per plant, 100 seed weight, percent seed set, plant height, days to 50 % flowering, stem diameter, head diameter and oil content in soybean. Pradhan et al. (2006) observed preponderance of non-additive gene action suggesting very good prospect for the exploitation of non-additive genetic variation for grain and its component characters in rice through hybrid breeding. The importance of non-additive gene for expression of different traits have also been reported by Ganesan et al. (1997), Ramalingam et al. (1997), Ganesan and Rangaswamy (1998), Bansal et al. (2000) and Thirumeni et al. (2000). The additive effects and gene interaction dominance x dominance (l) or other type of digenic complementary gene interaction can be exploited effectively by selection for improvement of the characters. Uses of reciprocal recurrent selection or bi-parental mating suggested improve the characters when both additive and non-additive gene
153
effects are involved in the expression of the traits. Presence of non-additive gene interactions in most of the plant characters as well as seed yield per plant and its contributing components indicated that conventional selection procedure may not be effective enough for improvement of seed yield in lentil. Therefore postponement of selection in later generations or intermating among the selected segregants followed by one or two generations of selfing could be suggested to break the undesirable linkage and allow the accumulation of favorable alleles for the improvement of this trait. The different types of gene effects estimated provided a test for gene action and are useful for analyzing the genetic architecture of a crop so as to further improve desirable traits. The estimates obtained from each cross may be unique to that cross and may not be applicable to the parental population. Additive genetic variance formed the major part of the genetic variance for the important seed yield component ‘100 seed weight’. Therefore, genetic improvement in seed yield per plant would be easier through indirect selection for a component trait such as the 100 seed weight than through direct selection for seed yield itself. Appropriate breeding methods such as reciprocal recurrent selection and biparental mating methods would be needed to improve the traits. These breeding strategies were useful for exploitation of both fixable and non-fixable types of gene action and were recommended by many breeders (Hayman 1958; Allard 1960; Falconer 1989; Kearsey and Pooni, 1996).
154
Chapter 6
Pattern of Inheritance of Qualitative Traits There are certain morphological traits that do not have economic value, but are important in practical plant breeding for identifying hybrids in a highly self-pollinated species. Seedling, flower and seed related colours are frequently used as marker characters in crossing programmes. Moreover, varietal identity, seed certification, roguing of off-type plants and genetic studies of other characters are some of the uses of these characters. In common lentil, purple and green are the two predominant hypocotyls colours. Among the agronomic traits, flowering time is one of the most important traits and wide variations exist in lentil. It is affected by the growing season and thus varieties are bred for specific geographical regions and seasons. The knowledge about different seedling/plant and seed related traits should serve to simplify the applied research, particularly the efforts to select parents for breeding, given the great difficulty in performing crosses. In the present studies, the pattern of inheritance for seedling growth habit, flower initiation, hypocotyls colour, seed coat pattern and cotyledon colour were investigated. Pattern of inheritance of seedling growth habit and flower initiation characters was studied in two crosses along with their respective reciprocals (Table 6.1). The seedling growth habits of female parents in both the crosses were erect and quick growth, and flower initiation was much early as compared to male parents who had spreading and slow growth at seedling stage, and also about 20-35 days later in flower initiation. The F1 plants of these crosses along with their respective reciprocals at seedling stage were spreading type and showed slow growth habit. Afterwards, all the plants were
155
observed late flowering. The F2 populations of these crosses segregated into plants with spreading habit and slow growth at seedling stage, and late in flower initiation with a good fit to 3: 1 ratio with nonsignificant χ2 values (χ2 = 0.00-3.03; P = 0.059-1.000). The F2 populations of the crosses (ILL 4605 x PL 406 & PL 406 x ILL 4605 and ILL 6821 x ILL 7715 & ILL 7715 x ILL 6821) segregated in a wider range of early and late flowering segregants which were far beyond the parental values in both directions generating a number of transgressive segregants for earliness as well as lateness but very few plants flowered like male parents which seemed a bottleneck to develop genotypes possessing desirable days to flower and seedling growth habit. Inheritance of hypocotyls colour was studied in cross ILL 5782 x ILL 2580 along with its reciprocal; the female parent having green hypocotyls colour while male parent (ILL 2580) had purple colour (Table 6.2). All the F1 plants had purple hypocotyls colour. The F2 population segregated with a good fit to the ratio of 3 purple: 1 green (χ2 = 0.132-1.169; P = 0.169-0.700). Seed coat pattern was studied in five crosses along with their respective reciprocals. The genotypes having no seed coat pattern (plain) were crossed as females with spotted males alog with their reciprocals. The seeds in F1 generation of all the crosses along with their reciprocals had spotted seed coat pattern (Table 6.3). The F2 populations in all the crosses segregated in to spotted and plain seed coat pattern with a good fit to 3: 1 ratio with nonsignificant χ2 values (χ2 = 0.005-0.469; P = 0.173-0.893). The pattern of inheritance in cotyledon colour was studied in two crosses along with their respective reciprocals. The genotypes having yellow cotyledon colour were crossed as females with red males and their reciprocals were also attempted. The seeds
156
Table 6.1: Segregation pattern and P values for seedling growth habit and flower initiation in lentil Cross F1 F2 Segregation P χ2 phenotype Female parent x Male parent Eqe Ssn Eqe x Ssn ILL 6821 x ILL 7715
All Ssn
130
409
0.247
0.840
ILL 4605 x PL 406
All Ssn
174
509
0.070
0.780
3.029 0.000
0.059 1.000
Ssn x Eqe ILL 7715 x ILL 6821 All Ssn 140 495 PL 406 x ILL 4605 All Ssn 149 447 Eqe: Seedlig erect, quick growth and early flower initiation Ssn: Seedling spreading, slow growth and normal flower initiation
157
Table 6.2: Segregation pattern and P values for hypocotyls colour lentil Cross F1 F2 Segregation χ2 phenotype Female parent x Male parent Purple Green
P
Green x Purple ILL 5782 x ILL 2580
All purple
489
158
0.132
0.700
Purple x Green ILL 2580 x ILL 5782
All purple
590
180
1.169
0.169
158
Table 6.3: Segregation pattern and P values for seed coat pattern in lentil Cross F1 F2 Segregation χ2 phenotype Female parent x Male parent Plain Spotted Plain x Spotted
P
ILL 5782 x ILL 2580 ILL 6468 x ILL 7556 ILL 6821 x ILL 7715 Masoor 93 x Turk Masoor ILL 4605 x PL 406
All spotted All spotted All spotted All spotted All spotted
76 65 59 77 59
209 200 184 253 155
0.469 0.005 0.022 0.145 0.156
0.475 0.893 0.725 0.464 0.453
Spotted x Plain ILL 2580 x ILL 5782 ILL 7556 x ILL 6468 ILL 7715 x ILL 6821 Turk Masoor x Turk Masoor PL 406 x ILL 4605
All spotted All spotted All spotted All spotted All spotted
77 55 67 35 49
247 192 195 112 151
0.263 1.055 0.020 0.145 0.027
0.636 0.173 0.893 0.720 0.881
159
were collected at maturity. It was observed that the crossed seeds possessed red cotyledon colour. The F1 generation was raised and analysis (Table 6.4) showed that all the plants were heterozygous (every plant had seeds with both red and yellow cotyledon colour) to cotyledon colour with a good fit to 3: 1 ratio with nonsignificant χ2 values (χ2 = 0.007-0.676; P = 0.444-0.940). In the F2 generation, plants segregated in to 1 (red cotyledon colour): 2 (red + yellow cotyledon colour): 1 (yellow cotyledon colour) ratio with nonsignificant χ2 values (χ2 = 0.022-2.323; P = 0.150-0.989). When the seeds of heterozygous plants harvested from F2 generation were counted, a good fit of 3 (red cotyledon colour): 1 (yellow cotyledon colour) ratio was observed with nonsignificant χ2 values.
Discussion Monogenic markers are useful for estimating the rate of crossing in predominantly self-pollinated crops (Senapathi and Roy, 1990). They also help in identification of F1 hybrids in the breeding programme. Heterozygous are not possible to detect in case of complete dominance for morphological markers. The results showed that spreading and slow growth habit, and late flowering characters controlled monogenically to erect and quick growth seedlings habit, and early flower initiation alleles as recessive. These three characters may be used as genetic markers in the inheritance studies. These characters have an importance also in hybridization where genotypes possessing erect and quick growth at seedling stage, and early flower initiation should be used as the female parent and the genotypes possessing spreading and slow growth at seedling stage, and late flowering as male parent. Crossed and selfed plants can be distinguished at the seedling stage and later on at flowering. The results of Emami and Sharma (1999) on growth habit, and Kant and Singh (1997), Tyagi and Sharma 160
Table 6.4: Segregation pattern and P values for cotyledon colour in lentil crosses Cross Segregation pattern
χ2
P
0.519
0.469
0.310
0.850
0.029
0.855
0.007
0.940
0.022
0.989
0.880
0.340
Female parent x Male parent Yellow x Red ILL 6468 x ILL 7556
F1 generation F1 plants harvested F2 Segregation
All plants heterozygous to cotyledon colour Red 2211 Yellow 760 Plants
Seeds (in F2 segregating plants) Red x Yellow ILL 7556 x ILL 6468
F1 generation F1 plants harvested F2 Segregation
Red Red +Yellow Yellow Red
73 143 67 6648
Yellow
2206
All plants heterozygous to cotyledon colour Red 2200 Yellow 731 Plants
Seeds (in F2 segregating plants)
Red Red +Yellow Yellow Red
69 139 70 6600
Yellow
2150 Contd…….
161
Cross
Segregation pattern
χ2
P
0.676
0.444
0.619
0.453
2.323
0.150
0.033
0.889
0.106
0.949
0.581
0.460
Female parent x Male parent Yellow x Red ILL 4605 x PL 406
F1 generation F1 plants harvested F2 Segregation
All plants heterozygous to cotyledon colour Red 1899 Yellow 657 Plants
Seeds (in F2 segregating plants) Red x Yellow PL 406 x ILL 4605
F1 generation F1 plants harvested F2 Segregation
Red Red +Yellow Yellow Red
59 101 49 5344
Yellow
1856
All plants heterozygous to cotyledon colour Red 1928 Yellow 637 Plants
Seeds (in F2 segregating plants)
Red Red +Yellow Yellow Red
51 105 54 1336
Yellow
464
162
(1989), Sharma et al. (1993) and Sarker et al. (1999a) on flowering also supported the present findings. Time to flowering plays a central role in determining the adaptation and productivity of the crop in short growing environments (Kumar and Abbo, 2001). The genetics of time to flowering needs to be sufficiently understood in order to fine-tune cultivars to the demands of a particular environment. Late flowering is dominant over earliness in chickpea (Gumber and Sarvjeet, 1996; Or et al. 1999; Kumar and van Rheenen, 2000; Anbessa et al. 2006). In short-season temperate environments, the duration of reproductive period is determined by the commencement of flowering and the end-of-season drought or frost that terminates seed setting and growth. A longer reproductive period, brought about by early-flowering alleles, could enhance seed yield in chickpea by allowing formation of a relatively large number of pods and through longer grain-filling duration (Or et al. 1999). Therefore, more progress could be made with respect to yield and earliness by incorporating the early flowering alleles into adapted genetic backgrounds. Constitutive earliness adapted to the prevailing environments can be considered as one of the best studied examples of effective escape from drought/heat constraints but the excessive earliness in the elite materials has often been correlated with constitutively low biomass and low grain yield potential (Reynolds et al. 2007). Initiation of flowering, in the segregating populations arose from the crosses between genotypes having less differences in flowering time, were observed well within the limits of the parents (data not presented) while in the population derived from the distant parents yielded transgressive segregations in flowering and maturity. Late flowering segregents did not perform well under our conditions as the temperature at pod/seed formation stage raised enough high and the pods produced immature seeds ultimately producing low seed yields. The other constraint observed to develop desirable
163
genotypes between the crosses of early and late flowering genotypes was that the early flowering recombinants had thin and less branchy plant type as was observed in wheat (Reynolds et al. 2007). Nevertheless, hybridization between macrosperma and microsperma types would leave tremendous scope for the improvement of lentil although early flowering/maturity trait is difficult to combine with high yield. The selection of early flowering segregants possessing spreading and slow growth habit under the prevalent climatic conditions is desirable and may prove fruitful for the improvement in lentil seed yield. Hypocotyls colour may be used as a genetic marker at seedling stage for the identification/inheritance studies. A single gene controlled this character. Purple hypocotyls colour was dominant to green. Same type of inheritance was observed by Verma and Nakhtore (1976), Ladizinsky (1979b) and Bukhsh and Iqbal, (2008). This trait has an importance in the hybridization programme where genotypes having green hypocotyls colour at seedling stage should be used as the female parent and the genotypes with purple colour as male parent. Crossed and selfed plants can be distinguished at the seedling stage. Ethnic preferences for specific seed coat colours and patterns of spotting influence the acceptance and marketability of lentil and hence, it is an important commercial attribute (Emami and Sharma, 2000). Present studies showed monogenic inheritance of seed coat pattern. The spotted seed coat was observed dominant over plain seed coat that are in confirmation to the studies of Ladizinsky (1979b) while the studies of Vandenberg and Slinkard (1990) showed that spotted and dotted seed coat patterns had codominant alleles, both of which were dominant to nonspotted seed coat pattern. They also reported that seed coat pattern was determined by a series of five alleles at another locus. This trait has significant importance in the hybridization programme. The
164
genotypes with plain seed coat pattern may be used as the female parent and the genotypes with spotted seed coat pattern as male parent. Crossed and selfed plants in the F1 populations can be distinguished at maturity before harvest. The cotyledon colours are commercially important attributes in Pakistan. Red (orange) colour is the predominant cotyledon colour in lentil. Monogeneic dominant red cotyledon colour over yellow in the present studies confirmed the findings of Bukhsh and Iqbal, (2008), Emami (1996b), Malaviya and Shukla (1990), Singh (1978), Sinha et al. (1987), Slinkard (1978), Tschermak-seysenegg (1928) and Wilson et al. (1970). This character has great importance in the conventional breeding. The genotypes with yellow cotyledon colour may be used as the female parent and the genotypes with red cotyledon colour as male parent. Crossed seeds can be distinguished at maturity before collection. The manifestation of red cotyledon colour to the crossed seeds suggested the use of cotyledon colour as a genetic marker in crosses to make sure that all the resulting seeds were hybrids. From the present studies, it may be concluded that an improved knowledge of the genetic basis of qualitative characters aimed to select new cultivars carrying such appealing traits will better support the lentil breeding programmes.
165
Chapter 7
Discussion and Conclusions The continuing need for improved crops to meet new environmental challenges and changing consumers’ demands create a constant requirement for genetic diversity, but the pool of natural diversity is shrinking with time largely because of the negative actions of humans (Guarino, 1999). The loss of genetic diversity commonly referred to as genetic erosion, results in the increasing vulnerability of crops to changing abiotic and biotic stresses and threatens global food security (Hawkes et al. 2000). Plant breeders and geneticists have addressed these stresses by identifying resistant/tolerant germplasm, determining the genetics involved and the genetic map positions of the resistant genes (Muehlbauer et al. 2006). In lentil, research efforts made through the introgression of genes from the ICARDA (International Center of Agricultural Research for the Dry Areas) germplasm to broaden the genetic base of exclusively pilosae lentil types with increased sensitivity to temperature limiting the flow of lentil germplasm into the IndoGangetic plain and consequently causing low yield potential in the region (Sarker and Erskine, 2002), have been culminated in the development of early, high yielding and disease resistant varieties in Bangladesh, India, Nepal and Pakistan (Chauhan and Singh, 1995b; Sarker et al. 1999b and 1999c; Sarker and Erskine, 2002; Sadiq et al. 2002 and 2006; Tufail et al. 1995). Despite the progress made in cultivar development in lentil, the released varieties could not make the way to boost up the yield per se and the average lentil yield in the region is still low as compared to other countries around the world. Further genetic progress demands more information on the inheritance of the key yield contributing traits and their associations with other plant traits according to the prevailing weather conditions of the target environment. Therefore, genetic studies are 166
important to answer specific questions relative to germplasm resources, types of genetic effects and modified selection methods to enhance cultivar development and upgrade the germplasm pools (Dudley and Moll, 1969; Moll and Stuber, 1974; Dudley, 1997). On average, around 50% of the productivity increases have been estimated to genetic improvement (Fehr, 1984). This study encompasses to the extent of genetic variation in the lentil germplasm, diversity among the parental lines used to develop segregating populations, estimations of nature of gene actions for seed yield and other important plant/yield related traits, and also the pattern of inheritance in some of the qualitative traits useful as morphological markers in conventional breeding programmes. Genetic variability of germplasm resources is of interest to plant breeders because it is necessary to sustain long-term genetic improvement of cultivars. The assessment of genetic variation and the behaviour of different genotypes in the germplasm with specific phenological responses provide the basis for adaptation to the climatic variables of the prevailing environment. The results obtained from germplasm evaluation applying different types of biometrical analyses (genotypic/phenotypic correlations, path analysis, multiple correlations, heritability and genetic advance estimates, and principal component analysis) have indicated the favourable importance of pods per plant, biomass per plant, 100 seed weight, plant height and productive branches while harvest index and days to flower showed contrasting effects in the improvement of lentil seed yield per plant. Summarized results presented in table 7.1 showed significant genetic variations of different plant and yield related traits in the germplasm indicating the scope and their warranty to use in the breeding programme (Tullu et al. 2001).
167
Table 7.1: Assessment of genetic variation through analysis of variance, correlations, path analysis and genetic parameters for important plant characteristics in lentil germplasm Parameter
Days to flower
Plant height
Primary branches plant-1
Secondary branches plant-1
Nodes on primary branch
Fertile nodes Nodes on on primary secondary branch branch
Fertile nodes on secondary branch
Pod length
Pod breadth
116.87 ± 16.34
37.24 ± 6.12
2.63 ± 0.19
18.95 ± 4.91
23.41 ± 4.01
12.71 ± 3.78
19.01 ± 3.47
9.45 ± 3.44
1.18 ± 0.17
0.62 ± 0.09
800.71 26.13 3.20 250.11**
112.33 35.51 6.58 17.06**
0.10 1.01 0.05 2.00**
72.56 6.69 2.77 26.19**
48.14 10.05 3.08 15.64**
42.92 6.81 1.20 35.72**
36.11 0.52 3.00 12.04**
35.55 2.84 1.43 24.89**
0.09 0.00 0.00 113.54**
0.03 0.00 0.00 77.19**
Days to flower
Genotypic Phenotypic
-0.1755 -0.1705
0.0425 0.0326
-0.0251 -0.0259
0.0529 0.0511
-0.3076* -0.3026**
0.0304 0.0286
-0.3868* -0.3783**
0.0034 0.0042
Plant height
Genotypic Phenotypic
1
0.1511 0.0928
0.0657 0.0669
0.3802* 0.3579**
0.1665 0.1596
0.3378* 0.3170**
0.2755* 0.2648**
Fertile nodes on secondary branch
Genotypic Phenotypic
0.2755* 0.2648**
0.2277* 0.1648
0.0280 0.0271
0.3077* 0.2849**
0.3957* 0.3778**
0.5122* 0.5337**
Pods plant-1
Genotypic Phenotypic Genotypic Phenotypic
0.2539* 0.2472* 0.0620 0.0596
0.1494 0.1091 0.0166 0.0128
0.2472* 0.2437* -0.2190* -0.2150*
0.0521 0.0460 -0.1446 -0.1382
0.4446* 0.4336** -0.2969 -0.2915**
Biomass plant-1
Genotypic Phenotypic
0.2094* 0.2056*
0.0625 0.0603
0.0808 0.0806
0.0101 0.0045
Harvest Index plant-1
Genotypic Phenotypic
0.2860* 0.2797**
0.1640 0.1321
0.1733 0.1695
0.1005 0.0910
Seeds per pod
Pods plant-1
100 seed weight
Biomass plant-1
Harvest Index plant-1
Seed yield plant-1
Analysis of variance Mean Standard deviation MS (genotype) MS (replication) MS (Error) F-ratio (genotype)
Correlations
100 seed weight
1.62 ± 0.22
168.25 ± 68.11
2.17 ± 0.84
14.69 ± 5.22
25.10 ± 7.05
3.82 ± 1.85
0.15 0.01 0.00 154.22**
13915.90 242.16 150.06 92.73**
2.11 0.02 0.01 336.18**
81.73 1.81 1.56 52.35**
149.28 18.82 1.70 87.72**
10.17 0.04 0.14 75.19**
0.2066* 0.0851
-0.1311 -0.1302
-0.2476* -0.2457*
-0.1639 -0.1632
-0.3852* -0.3825**
0.0797 0.0798
0.0707 0.0386
0.0304 0.0297
0.2540* 0.2472*
0.0620 0.0596
0.2094* 0.2056*
0.2860* 0.2797**
-0.0558 -0.0554
1.0000 1.0000
-0.1615 -0.1579
-0.5192* -0.1854
0.0631 0.0618
0.2885* 0.2982**
-0.1265 -0.1305
0.2116* 0.2158*
0.1773 0.1836
0.1688 0.1666
0.0689 0.0898 -0.0692 -0.0755
0.2885* 0.2982** -0.1265 -0.1305
-0.3128* -0.3091** 0.6009* 0.5971**
-0.7276* -0.2789** 0.3577* 0.1378
0.0999 0.0992 -0.0842 -0.0841
1.0000 1.0000 -0.1267 -0.1298
-0.1267 -0.1298 1.0000 1.0000
0.7418* 0.7389** 0.4302* 0.4256**
0.6479* 0.6448** 0.3562* 0.3511**
0.4826* 0.4791** 0.3130* 0.3102**
0.2350* 0.2270*
0.0258 0.0380
0.2116* 0.2158*
0.0451 0.0443
-0.3470* -0.1317
0.2372* 0.2367*
0.7418* 0.7389**
0.4302* 0.4256**
1.0000 1.0000
0.8332* 0.8331**
0.7078* 0.7030**
0.1232 0.1169
0.1442 0.1528
0.1773 0.1836
0.0583 0.0576
-0.3659* -0.1370
0.1758 0.1754
0.6479* 0.6448**
0.3562* 0.3511**
0.8332* 0.8331**
1.0000 1.0000
0.2318* 0.2280*
-0.1165 -0.1153
-0.5214* -0.5184**
Path analysis Direct effect
Indirect effect
rg with seed yield plant
Plant height Secondary branches plant-1 Fertile nodes on primary branch Fertile nodes on secondary branch Pods plant-1 100 seed weight Biomass plant-1 Harvest Index plant-1
–0.133 0.114
1 –0.0087
–0.0177 –0.0259
0.0075 1
–0.0097 –0.0020
–0.131 0.185 0.249 0.160 1.437 –1.091
–0.0219 –0.0566
0.0007 –0.0001
0.0509 0.0052
–0.0129 0.0491
–0.0221
0.0065
0.0491
–0.0365
–0.0267
0.0032
–0.0337 –0.0082 –0.0278 –0.0379
–0.0175 –0.0019 –0.0073 –0.0193
0.0281 0.0249 0.0092 0.0197
0.0131 –0.0963
0.0087 –0.0006
0.0631 0.0615
0.0099 –0.0351
0.3009 0.0898
–0.3120 –0.1890
–0.0557 –0.0681
–0.0109
1
0.0003
0.0731
–0.0079
–0.0519
0.0010
1
–0.0637
0.0542
0.0010
0.1105
–0.0476
0.3377
–0.1344
0.2489
0.0262
–0.0964
0.0018
0.0717
–0.0203
0.3041
–0.1934
0.1688
–0.0013 0.0037 –0.0003 –0.0026
–0.0584 0.0389 –0.0309 –0.0162
0.0001 –0.0001 0.0391 0.0003
0.0533 –0.0233 0.0391 0.0327
0.0508 –0.0976 –0.0073 –0.0095
–0.1351 0.0664 –0.0645 –0.0679
0.0029 –0.0024 0.0068 0.0050
1 –0.0315 0.1844 0.1611
–0.0203 1 0.0689 0.0571
1.0660 0.6183 1 1.1974
–0.7068 –0.3886 –0.9091 1
0.4826 0.3129 0.708 0.2318
Genetic parameters Phenotypic variance
37.44
0.04
24.19
16.05
14.31
12.04
11.85
0.03
0.04
0.05
4638.63
0.70
3.39
49.76
27.24
Genotypic 265.84 35.25 variance PCV 2.28 1.01 GCV 2.27 0.95 2 99.60 94.15 h GA (% of mean) 28.68 31.87 ** and *: Significant at P < 0.01 and P < 0.05
266.91
0.02
23.26
15.02
13.91
11.04
11.37
0.03
0.01
0.05
4588.61
0.70
3.34
49.19
26.72
0.01 0.07 50.00 7.22
1.28 1.23 96.16 51.48
0.69 0.64 93.58 32.98
1.13 1.09 97.20 59.56
0.63 0.58 91.69 34.46
1.25 1.20 95.94 72.06
0.02 0.02 99.20 29.66
0.06 0.01 25.00 9.38
0.03 0.03 99.40 27.78
27.57 27.27 98.92 82.49
0.32 0.32 99.30 79.26
0.89 0.89 98.53 98.94
1.98 1.96 98.85 56.51
1.85 1.82 98.09 71.82
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-1
Phenology has been observed the most important single factor influencing genotypic adaptation in annual crops (Richards, 1991; Hamblin, 1994). In this study, flowering time in the germplasm was observed as a key limiting factor. Most of the genotypes were observed late to flower, and due to more rains during late in the growing period (Table 3.2) pod formation continued but the genotypes could not cope with the increasing temperature (at seed development stage) at harvest producing immature seeds with low seed weight and yield. Low seed weight and yield have been hypothesized in peas due to competition between sinks for available photosynthates (McPhee and Muehlbauer, 2001) and higher temperatures resulted in flower and pod abortion by reduced seed-filling period (Nielson, 2001). Late flowering lentil accessions have been observed to develop larger biomass but with low harvest index (Erskine et al. 1989) as was observed in this study. Seed yield improvements have been suggested by using accessions with high straw production while maintaining harvest indices similar to current estimates in pea (McPhee and Muehlbauer, 2001). The consequences of high rainfall during flowering and pod/seed development stages in this study were observed through direct negative contribution of harvest index and positive direct effect of biomass with seed yield. Several studies conducted on different crop plants including lentil have revealed the importance of flowering time well suited to the specific environments and adverse effects of high temperature at pod development and seed set stages (chapter 3). On the other hand, beneficial associations of yield contributing characters such as biomass, pods per plant, 100 seed weight and fertile nodes on secondary branches along with their direct effects on seed yield except for harvest index indicate the possibility for improved selection based on these traits if the flowering of the genotypes coincides with the climate of the growing environment. Estimates of higher heritability coupled with higher genetic advance observed for yield contributing traits such as biomass per plant, pods per plant, 100 seed weight, and fertile nodes on secondary branch also
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indicate the chance for improvements through direct selection in these traits keeping synchronized phenology of the genotypes. Selections made on the basis of yield contributing characters in soybean have been observed to play direct roles in yield improvements (Cui and Yu, 2005). Lentil being indeterminate plant type suffers problems of crop durations, seasonal cumulative rainfall as well as its distribution throughout the growing season, and temperature fluctuations specifically at flowering and pod setting stages causing low seed yield. These studies also indicated the needs for selection of genotypes having early phenology with determinate growth habit evident from the contrasting associations of days to flowering with seed yield per plant and, also with fertile nodes on primary and secondary branches, pods and biomass per plant. Under excessive moist conditions, lentil plants grow further with the formation of more pods at the apical parts, and with extended flowering period they continue to develop flowers and pods due to its indeterminate growth habit (Sarker et al. 2003) but abrupt rise in temperature during pod-filling stage along with low relative humidity (less than 50% during the months of March and April) hastened pods with fully-developed seeds resulting in low seed weight and yield per plant. Often the length of the growth period and environmental conditions have been observed of paramount importance in matching the cultivar’s needs with its climate to optimize yield (Friend, 1985), and largest reductions have been projected in southern crop areas of the US and Canada due to increased temperatures and reduced water availability. For which emphasis has been given to the development of proactive mitigation measures (a proactive measure focuses on preparations in advance of the event to help mitigate the impact on the system) to cope with the weather variability/changes and preserve the agricultural systems (Motha and Baier, 2005). The adverse effects of high temperature and low humidity on pod setting, forced maturity coupled with limited branching and restricted dry matter production, and
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seed yield in lentil have been observed by several scientists (Chandra and Asthana, 1988; Sarker et al. 2003; Summerfield, 1981 and Summerfield et al. 1989; Muehlbauer et al. 2006). To achieve higher seed yields coupled with increases in seed size, it would be imperative to develop cultivars which can partition greater amounts of photosynthates to the seed late in the season. Such type of adaptations has also been suggested in peas (McPhee and Muehlbauer, 2001). In other words, an increase in vegetative biomass prior to flowering, rapid crop growth and higher dry matter partitioning to seed combined with early flowering and maturity may prove beneficial during selection while developing improved varieties. The selection of genotypes, with determinate growth habit (in which flowering period is condensed and a greater proportion of nodes bear pods, the assimilates may be partitioned to pods rather than to continued vegetative growth) may help counteract with the contrasting effects of harvest index. Low crop harvest indices in wheat have been observed attributable to poor conservation of light, water and nutrients to grain potential yields (Riffkin and McNeil, 2006). Selection towards early phenology of the genotypes coupled with early and rapid biomass development for the realization of improved seed yields in lentil has been emphasized. So, it is likely to select genotypes with shorter vegetative and generative growth periods along with higher assimilate partitioning to seeds in those regions with little or no rainfall during later stages of the crop growth and development. For high rainfall coupled with low temperatures during reproductive phases of the plants, longer vegetative and generative growth periods, high response to rainfall and more favourable photosynthetic partitioning genotypes may prove useful (Kusmenoglu and Muehlbauer, 1998; Berger, 2006; Andrews and McKenzie, 2007). Present studies revealed that seed yield per plant may be enhanced through the indirect selection of tall bushy plants bearing more productive branches provided the phenology (negative associations of pods per plant with days to
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flower) of the plants synchronized with the growing climatic conditions which is self explanatory from the favouable associations of plant height with nodes on primary and secondary branches, and also with fertile nodes on secondary branches. The higher heritability estimates of pods per plant and its favouring associations with plant height, secondary branches, and fertile nodes on primary and secondary branches but negative correlation with days to flower also support the importance of phenology and productive bushy plants in integrating the genotypic variation towards selection. It means that tall bushy plants may tend to produce more nodes on the primary branches which in turn develop more secondary branches productive thus contributing ultimately to improved seed yield per plant. This may be achieved by selecting the genotypes which are able to partition the photosynthetic assimilates to the seeds at harvest because additional photosynthetic capacity from a more leafy canopy is available to meet the increased demands for carbon assimilate to dry matter accumulation (Silim et al. 1993). For this, effective crop duration, resource capture and dry matter accumulation may favour to higher biomass and improved seed yield by the genotypes well-adapted to the growing environment (Whitehead et al. 2000). Seed yield per plant, arguably the most important trait, a polygenic in nature, difficult to improve, and highly influenced by the environment, may be improved through indirect selection of seed yield contributing traits with the restriction that other characters may not suffer and the phenology of plants may suit to the growing environment. Characters such as plant height, productive branches, biomass and harvest index have been observed important for the improvement along with phenology (days to flowering) and seed size (Table 7.1). The importance of different characters in lentil has been emphasized in different studies according to the nature of the material evaluated and the environment where the plants were grown. However, morphological traits representing the action of
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numerous genes containing high information value can be unreliable owing to a strong influence of the environment (Smýkal et al. 2008). Besides, it is very often that selection for yield through yield components is limited by negative correlations among yield components i.e. yield compensation (Adams, 1967). Therefore, it is important to follow an appropriate strategy for the improvements during selection and the use of parents for production of segeregating populations according to the prevalent weather conditions of the specific region. Molecular and biochemical markers have revealed that lentil has relatively low levels of genetic variation in comparison to other plant species (Alvarez et al. 1997; Durán and Pérez de la Vega, 2004; Eujayl et al. 1997; Ferguson, 1998c; Ford et al. 1997; Sonante and Pignone, 2001). High estimates of similarity indices using RAPD markers indicated relatively low genetic diversity among the parental lines. Despite this, the present RAPD analysis was informative enough for differentiation among the lentil parental lines. No clear lineage was observed in clusters using both morphological and molecular markers regarding relationship between geographic pattern and genetic diversity as well as with visible characters of the parental lines whereas geographical separation with physical barriers and genetic barriers to crossablity is believed to give rise to genetic diversity among genetic materials (Singh, 1990). No definite correspondence between geographic origin and genetic diversity has been observed in other crops like field pea (Gemechu et al. 2005) and safflower (Khan et al. 2008) suggesting that parental selection should be made on the basis of systematic assessment of genetic distance in a specific population rather than on geographic difference. However, whether differences in geographic origin (source) necessarily imply genetic distance in parental selection for hybridization is still a matter of some controversy. Nevertheless, geographic diversity may serve as an index of genetic diversity in parental selection (Joshi and Dhawan, 1966) while others argued that genetic
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divergence was not apparently related to geographic diversity in some crops (Rezai and Frey, 1990; Katule et al. 1992). Selection of parents may be approached in two ways: a priori and a posteriori choice (Baenziger and Peterson, 1992). The a priori choice consists of selection methods based on per se parent performance, such as midparental value, divergence according to coefficient of parentage, character complementation, multivariate analysis and parental distances, least squares, parental complementation, and ideal genotype. In the a posteriori choice, parents are evaluated on the basis of F1, F2 data and advanced generations. A long period of time is necessary to choose parents in this way. The a priori choice is based on the fact that heterosis is a relative measure of two generations - the parental and the progeny, and was initially made on the basis of morphological descriptors and further reinforced by data from molecular markers. This choice assumes that the divergence between any two parents expresses the allelic differences between them (Dias et al. 2003). The genotypes grouped into the same cluster presumably diverge very little from one another. Crossing of genotypes belonging to the same cluster is not expected to yield desirable segregants. Consequently, a crossing programme should be formulated such that the putative parents belonging to different characters. Therefore, crosses between the members of clusters separated by inter-cluster distances are likely seemed to be beneficial for further improvement. Significant differences among the parental lines (chapter 4) as well as among the generations (chapter 5) developed from these parental lines, for different plant traits indicated variations among the parental lines favourable for their use in the breeding programme. Several reports in the literature (Chahal and Gosal, 2002; Chandel and Joshi, 1983; Gemechu et al. 1997) have indicated that crosses between parents with maximum divergence would be more responsive to improvement since they were likely to produce higher heterosis and desirable genetic recombination, and segregation in their progenies
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may help to develop varieties with broad genetic base raising the yield ceilings. On the other hand, there are several causes like epistasis (Boppenmaier et al. 1992), genotypeenvironment interactions (Dias et al. 2003), lack of linkage disequilibrium (Charcosset et al. 1991), lack of linkage between genes controlling the traits and the markers used (Bernardo, 1992), gene pool with a narrow genetic base (Maroof et al. 1997), increased genetic similarity in a gene pool due to strong selection pressure (Barbosa et al. 2003), differences in the contributions of the marked DNA regions (Kwon et al. 2002a & b) and high degree of improvement of the gene pool used (Dias et al. 2003) that suggest non-linear relationships of divergence and heterosis (Sant et al. 1999). In the present studies, lower mean values for majority of the characters in F1 and/or F2 generations of different crosses (Appendices XXIII-XXXIV) than the respective better parents indicated lack of heterosis and significant mean effect (m) in all the crosses as well as in characters (Tables 5.2-5.18). This may be attributable to one or more of the causes for the nonlinear diversity-heterosis relationship described above. While higher estimates of dominance effect (h) than additive effect (d) indicated that the parents used in the present studies were distant for the concerned traits and epistatic interactions of additive x additive (i), additive x dominance (j) and dominance x dominance (l) seemed integral components of the genetic formation of these traits. In crop improvement, lower seed yield of most progenies from a cross between any two elite parents of the same species or subspecies suggested the important role of epistasis for seed yield and its components (Li et al. 1997). It has also been observed in peanut that the less divergent parental lines are more likely to play additive gene effects in inheritance of quantitative traits (Isleib and Wynne, 1983) and as the diversity between parents increases, dominance effects and epistatic variations have significant roles in the inheritance of quantitative traits (Halward and Wynne, 1991) which may have implications in choosing an appropriate selection strategy. For a trait governed by multiple and quantitative trait loci
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(QTL) and/or co-dominantly inherited genes, a more holistic genome mapping approach to identify genomic locations, interaction and subsequent molecular markers for accurate trait selection (Muehlbauer et al. 2006) may prove fruitful. Using association mapping, entire genomes can be scanned for markers associated with qualitative and quantitative traits. The association mapping approach may allow plant breeders to break out of restrictive F1derived mapping populations and employ any plant population including those from breeding programmes or germplasm collections to conduct marker-trait association studies (Flint-Garcia et al. 2003). Parents selected on the basis of contrasting qualitative plant traits may contribute to speed up the crossing work during conventional breeding. Plant habit and the associated trait of crop height have been observed essential agronomic characteristics because of their influence on competition for light between neighbours. The monogenic genetic control (dominant) in the present inheritance studies of qualitative characters (growth habit, hypocotyls colour, seed coat pattern and cotyledon colour) has indicated their easier use in the lentil improvement programmes. Many simply inherited traits have been placed on lentil genetic maps. By knowing the map position of a gene, the presence of the gene can be diagnosed using flanking DNA markers without waiting for the gene effect to be present in the phenotype (Paterson et al. 1991). In the crosses of more distant parents in flower initiation showed transgressive segregations and late flowering segregants did not perform well under our conditions due to high temperature at pod/seed formation stage producing pods with immature seeds (low seed weight) ultimately low seed yield. The same trend was also observed during germplasm evaluation where late flowering genotypes did not show better performance producing high biomass with low harvest index. Besides, extra early flowering recombinants had thin and less branchy plant type and produced lower seed yield, not fit well under the prevailing conditions. Therefore, selection of recombinants possessing spreading/semi erect growth habit with appropriate phenology suited to the
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environmental variables seemed to be fruitful. This is a demending task for lentil improvement due to the non-availability of frequent occurrence of recombinants having such type of combinations of these characters during selection. Epistasis gene interactions in each cross for different plant traits have been found different from each other. Among the epistasis interactions, prevalence of larger effects of dominance x dominance (l) interaction than either additive x additive (i) or additive x dominance (j) interaction in majority of the crosses for days to flower and mature indicated predominant duplicate type of gene interactions for these traits that was also observed in germplasm evaluation studies for days to flower in chapter 3 (higher estimates of h2 with low genetic advance). Thus, selection for increased trait values should be more efficient when it is practiced on individual allelic combinations at two or more loci. If so, gene/QTL mapping needs to place more emphasis on identifying the best multilocus gene combinations (Li et al. 1997). During evaluation of germplasm (chapter 3) and also in the segregating populations (chapter 6), long duration (late flowering) genotypes with short reproductive phase could not perform well due to the substantial high temperatures late during the growing season. Therefore, selection for longer duration genotypes seems to be beneficial where frequent moisture and mild temperatures are prevalent, and vice versa. In other words, selections with appropriate values of phenology (flowering and maturity) which are well suited to the prevailing growing environments may perform better. Relatively longer and cooler season has been observed beneficial for obtaining higher seed yields in lentil (Naurai, 1991). In grain legume crops including lentil, genotype x environment interactions have been described for different plant/yield related traits indicating the importance of selection for targeted environment(s). Thus, while selecting the exotic germplasm lines for inclusion in the breeding programme, it is important to consider the genetic background and agronomic performance of the lines, as it will be useful in
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predicting its behavior in hybrid combinations with the adapted genotypes (Upadhyaya et al. 2007). Duplicate type of gene interactions for seed yield per plant and its contributing traits exhibiting complex nature of inheritance may tend to hinder the progress making it difficult to fix the genes at increased level of manifestation. Although higher heritabilty and genetic advance estimates suggested additive type of gene actions for seed yield per plant, biomass per plant, harvest index per plant, pods per plant and seed weight in the germplasm evaluation studies (chapter 3) but the prevalence of non-additive gene interactions (chapter 5) in most of the plant characters as well as seed yield per plant and its contributing components (attributable due to the differences in the germplasm material and the segregating populations as well as the differences in weather conditions during the years of evaluation) indicated that conventional selection procedure may not be effective enough for improvement of seed yield in lentil. Thus, the integrations of different types of breeding strategies like postponement of selection till further generations, intermating among the selected recombinants for the breakage of undesirable linkages and accumulation of favourabe alleles in later generations, hybrid breeding and reciprocal recurrent selections may prove helpful for the improvements to the respective crosses in parallel to the nature of inheritance for specific traits. The use of molecular markers along with quantitative traits loci (QTLs) mapping for different morpho-agronomic plant traits may contribute in the lentil improvements.
Conclusions •
Utilization of diversified germaplsm possessing different desirable plant traits may prove helpful by incorporating these traits in the lentil improvement process.
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•
Flowering time in the germplasm as well as in the segregating populations was observed as a key limiting factor.
•
Selection of late flowering genotypes may prove favouable for long crop durations where high rainfalls and low/mild temperatures are prevalent late in the growing season and vice versa.
•
Selection for tall bushy plants possessing more productive branches may support to acquire higher biomass and produce higher seed yield provided the phenology (flowering) of the plants synchronized with the prevailing environment, and improved harvest index through proper utilization of radiation intercept, and balanced source and sink relationships of photothsynthates.
•
Improvements in harvest index seem very demanding task.
•
Relatively low genetic diversity was observed among the parental lines through RAPD technique.
•
Generally, no definite correspondence between geographic pattern and genetic diversity, and visible characters was observed in parents through morphological as well as molecular markers.
•
Inheritance studies on qualitative characters may prove to be effectively beneficial for the identification of crossed seeds in conventional breeding and for selection of improved recombinants maintaining the consumers’ demand especially spotted seed coat pattern and red cotyledon colour.
•
Dominance effect (h) was found to be more significant than the additive effect (d) in most crosses. The epistatic interactions (i, j and l) also played significant roles in controlling the inheritance of different traits. Additive and non-additive gene interactions (additive, dominance, and epistasis) on seed yield per plant and its contributing traits suggested that appropriate selection procedures may be adopted to
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improve the traits accordingly (depending upon the mode of inheritance in the respective traits and crosses, and the growing environments). •
The integration of classical plant breeding with molecular markers and QTLs mapping for genetic studies of different traits of interest also seemed beneficial to assist in the lentil improvement.
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208
Appendix I: Details of parental lines used for the production of segregating populations in the inheritance studies S. No.
Genotype
Pedigree
Source
1
ILL 4605
Precoz
Argentina,
Erect and quick seedling growth, early maturing, short stature, Large seed size, yellow cotyledon colour, non-spotted seed coat
2
ILL 5782
ILL 883 x ILL 470
ICARDA, Syria
Normal maturing, short stature, large seed size, red cotyledon colour, non-spotted seed coat
3
ILL 6024
ILL x ILL 3458
ICARDA, Syria
Normal maturing, short stature, large seed size, red cotyledon colour, non-spotted seed coat
4
Pant L 406
Selection from P495
India
Normal maturing, tall, small seed size, red cotyledon colour, spotted seed coat
5
ILL 6821
ILL x ILL 4605
ICARDA, Syria
Erect and quick seedling growth, early maturing, short stature, large seed size, red cotyledon colour, non-spotted seed coat
6
ILL 8117
ILL 6243 x ILL 5883
ICARDA
Normal maturing, tall, small seed size, red cotyledon colour, non-spotted seed coat
7
ILL 7556
GL – I
India
Normal maturing, tall, small seed size, red cotyledon colour, spotted seed coat
8
ILL 7715
PL 81-17
India
Normal maturing, tall, small seed size, red cotyledon colour, spotted seed coat
9
ILL 6468
ILL 5562 x ILL 2501
ICARDA, Syria,
Normal maturing, Short stature, small seed size, yellow cotyledon colour, non-spotted seed coat
10
ILL 2580
-
India
Normal maturing, tall, small seed size, red cotyledon colour, spotted seed coat
11
Masoor 93
ILL 4400 x L 18-12
Pakistan
Normal maturing, tall, small seed size, red cotyledon colour, non-spotted seed coat
12
Turk Masoor
Bulk sample from local market
-
Salient Features
Selection from bulk sample of the market. Flowering starts early but flowers die, Early maturing, short stature, large seed size, red cotyledon colour, spotted seed coat
209
Appendix II: Sequence of primers used for genetic diversity in lentil No. Name Sequence 1
OPA-1
5/-CAGGCCCTTC-3/
2
OPA -13
5/-CAGCACCCAC-3/
3
OPB-1
5/-GTTTCGCTCC-3/
4
OPB-5
5/-TGCGCCCTTC-3/
5
OPB-9
5/-TGGGGGACTC-3/
6
OPB-19
5/-ACCCCCGAAG-3/
7
OPN-2
5/-ACCAGGGGCA-3/
8
OPN-4
5/-GACCGACCCA-3/
9
OPN-5
5/-ACTGAACGCC-3/
10
OPN-6
5/-GAGACGCACA-3/
11
OPN-13
5/-AGCGTCACTC-3/
12
OPN-14
5/-TCGTGCGGGT-3/
13
OPN-15
5/-CAGCGACTGT-3/
14
OPN-16
5/-AAGCGACCTG-3/
15
OPN-18
5/-GGTGAGGTCA-3/
210
Appendix-III: Meteorological data for the growing season of lentil during 2001-02 Air Temperature (oC) Maximum Minimum
Month
Relative Humidity (%)
Total Rain Fall High Low Mean High Low Mean High Mean Low Mean (mm) November 31.8 25.2 28.3 21.0 08.5 11.9 94 78.3 30 46.9 03.2 December
27.0
13.0
22.4
11.4
04.5
07.4
100
89.6
31
55.3
T
January
25.0
13.2
19.8
14.0
01.0
04.1
100
87.7
25
49.2
T
February
26.0
15.7
22.4
15.0
01.5
07.7
100
82.2
23
43.0
01.0
March
35.5
18.0
29.6
21.1
08.8
13.8
94
76.4
22
42.2
09.5
April
43.5
26.1
35.4
28.6
14.4
20.8
90
53.1
13
33.4
05.2
T: Traces Source: Plant Physiology Section, Ayub Agricultural Research Institute, Faislalabad
211
Appendix IV: Mean square values for days to flower in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Error
Total
1
10
17
Ps, F1s vs BCs, F2s
Degree of Freedom
2
5
1
1
ILL 5782 x ILL 2580
2.073
19.047**
2.870
29.007**
13.202**
0.088
50.067**
0.666
6.238
ILL 2580 x ILL 5782
1.097
18.743**
2.870
28.754**
1.984
25.134**
34.973**
1.652
6.614
ILL 6024 x ILL 8117
1.400
18.760**
0.810
41.710**
22.040**
2.030
27.210**
0.830
6.170
LL 8117 x ILL6024
1.388
19.837**
0.807
40.500**
30.150**
5.260*
22.467**
0.732
6.428
ILL 6468 x ILL 7556
0.685
9.910**
2.535*
0.080
44.010**
1.439
1.485
0.462
3.267
ILL 7556 x ILL 6468
0.008
9.812**
2.535*
0.001
36.260**
5.152**
5.109**
0.405
3.125
ILL 6821 x ILL 7715
3.220
428.170**
1153.710**
602.050**
164.330**
14.400
206.380**
3.840
128.570
ILL 7715 x ILL 6821
0.510
433.640**
1153.710**
590.530**
196.650**
57.210**
170.080**
4.650
130.330
Masoor 93 x Turk Masoor
0.834
149.649**
480.615**
115.520**
1.127
2.376
148.609**
1.332
44.896
Turk Masoor x Masoor 93
0.718
147.566**
480.615**
110.014**
2.100
0.031
145.067**
1.226
44.208
ILL 4605 x PL 406
7.400
346.260**
384.000**
1192.350**
10.010
66.090**
78.880**
4.200
105.180
PL 406 x ILL 4605
4.960
343.640**
384.000**
1171.280**
77.760**
29.290**
55.900**
1.360
102.460
** and *: Significant at P < 0.01 and P < 0.05 respectively
212
1
1
Appendix V: Mean square values for days to maturity in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Ps, F1s
Error
Total
vs BCs, F2s Degree of Freedom
2
5
1
1
1
1
1
10
17
ILL 5782 x ILL 2580
1.420
18.480**
31.740**
6.480
0.844
28.250**
25.087**
1.359
6.402
ILL 2580 x ILL 5782
1.760
18.527**
31.740**
7.347
0.375
28.577**
24.593**
1.404
6.482
ILL 6024 x ILL 8117
3.113
38.164**
114.407**
40.801**
0.002
29.954**
5.656
1.527
12.489
LL 8117 x ILL6024
2.367
36.380**
114.407**
35.001**
0.107
25.063**
7.322*
0.887
11.500
ILL 6468 x ILL 7556
2.758
17.750**
3.760
67.860**
4.167
5.645*
7.296*
0.973
6.116
ILL 7556 x ILL 6468
1.703
15.278**
3.760
54.254**
2.100
5.634*
10.641**
1.020
5.294
ILL 6821 x ILL 7715
4.459
72.946**
260.700**
58.861**
28.167**
12.268**
4.733
1.040
22.591
ILL 7715 x ILL 6821
2.990
78.323**
260.700**
62.533**
44.282**
14.258**
9.842**
0.910
23.923
Masoor 93 x Turk Masoor
0.044
208.124**
316.827**
201.336**
384.000**
89.780**
48.676**
0.380
61.441
Turk Masoor x Masoor 93
0.013
206.480**
316.830**
213.560**
380.010**
80.349**
41.678**
0.566
61.065
ILL 4605 x PL 406
1.370
199.570**
535.815**
456.020**
2.220
0.190
3.610
1.700
59.860
PL 406 x ILL 4605
4.412
196.592**
535.815**
439.067**
5.607
0.752
1.717
1.876
59.444
** and *: Significant at P < 0.01 and P < 0.05 respectively
213
Appendix VI: Mean square values for plant height in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Ps, F1s
Error
Total
10
17
vs BCs, F2s Degree of Freedom
2
5
1
1
1
1
1
ILL 5782 x ILL 2580
1.232
8.916**
15.844**
2.531
3.082
0.989
22.133**
0.727
3.195
ILL 2580 x ILL 5782
0.436
22.870**
15.844**
1.004
3.375
11.907*
82.219*
1.122
7.438
ILL 6024 x ILL 8117
0.588
100.855**
361.849**
65.342**
49.307**
0.088
27.689**
1.055
30.353
LL 8117 x ILL6024
0.921
109.787**
361.849**
61.587**
46.760**
7.157**
71.581**
0.476
32.678
ILL 6468 x ILL 7556
0.545
58.672**
182.050**
46.561**
10.800**
0.524
53.423**
0.735
17.753
ILL 7556 x ILL 6468
5.360
52.669**
182.050**
30.031*
8.050
15.587
27.627
6.013
19.659
ILL 6821 x ILL 7715
0.591
107.629**
72.107**
434.142**
17.682*
12.467
1.748
2.558
33.230
ILL 7715 x ILL 6821
0.274
98.444**
72.107**
373.556**
16.335*
30.161**
0.061
2.056
30.196
Masoor 93 x Turk Masoor
1.763
57.361**
19.260**
106.823**
8.760
5.814
146.148**
2.004
18.257
Turk Masoor x Masoor 93
0.971
43.875**
19.260**
116.790**
1.170
1.042
81.110**
0.649
13.401
ILL 4605 x PL 406
2.213
59.076**
44.554**
58.140**
0.094
0.057
192.530**
0.774
18.091
PL 406 x ILL 4605
1.180
63.845**
44.554**
77.501**
0.770
1.566*
194.834**
0.296
19.091
** and *: Significant at P < 0.01 and P < 0.05 respectively
214
Appendix VII: Mean square values for primary branches in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Error
Total
1
10
17
Ps, F1s vs BCs, F2s
Degree of Freedom
2
5
1
1
1
ILL 5782 x ILL 2580
0.026
0.116**
0.282**
0.094**
0.034
0.101**
0.067*
0.007
0.041
ILL 2580 x ILL 5782
0.026
0.116**
0.282**
0.094**
0.034
0.101**
0.067*
0.007
0.041
ILL 6024 x ILL 8117
0.057
0.371**
0.019
0.022
1.550**
0.262**
0.001
0.009
0.121
LL 8117 x ILL6024
0.001
0.316**
0.019
0.039**
1.307**
0.205**
0.011
0.005
0.096
ILL 6468 x ILL 7556
0.003
0.155**
0.220**
0.101*
0.007
0.192**
0.252**
0.017
0.056
ILL 7556 x ILL 6468
0.017
0.082*
0.220**
0.023
0.000
0.026
0.137**
0.019
0.037
ILL 6821 x ILL 7715
0.045
0.332**
0.540**
0.436**
0.304*
0.307*
0.073
0.037
0.124
ILL 7715 x ILL 6821
0.006
0.409**
0.540**
0.642**
0.184
0.669**
0.008
0.018
0.131
Masoor 93 x Turk Masoor
0.012
0.068**
0.000
0.073**
0.007
0.213**
0.048*
0.009
0.027
Turk Masoor x Masoor 93
0.002
0.021
0.000
0.073**
0.027
0.003
0.002
0.012
0.013
ILL 4605 x PL 406
0.004
0.154**
0.002
0.005
0.002
0.026
0.736**
0.021
0.058
PL 406 x ILL 4605
0.036
0.188**
0.002
0.027
0.010
0.005
0.898**
0.020
0.072
** and *: Significant at P < 0.01 and P < 0.05 respectively
215
1
Appendix VIII: Mean square values for secondary branches in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Ps, F1s
Error
Total
10
17
vs BCs, F2s Degree of Freedom
2
5
1
1
1
1
1
ILL 5782 x ILL 2580
5.985*
36.264**
123.307**
40.201**
0.050
16.956**
0.806
1.272
12.118
ILL 2580 x ILL 5782
2.168
53.295**
123.307**
27.134**
0.735
105.997**
9.302**
0.512
16.231
ILL 6024 x ILL 8117
1.787
29.509**
66.633**
7.088
49.594**
5.412
18.819**
1.833
9.968
LL 8117 x ILL6024
1.759
38.932**
66.633**
2.957
57.350**
23.052**
44.667**
1.818
12.727
ILL 6468 x ILL 7556
0.614
5.085*
7.370*
0.211
7.594*
2.486
7.762*
1.248
2.302
ILL 7556 x ILL 6468
0.080
5.033*
7.370*
0.957
4.335
3.042
9.461*
1.196
2.193
ILL 6821 x ILL 7715
0.225
19.450**
1.760
77.917**
1.760
15.513**
0.302
0.591
6.095
ILL 7715 x ILL 6821
0.297
13.529**
1.760
49.501**
0.082
15.420**
0.880
0.412
4.257
Masoor 93 x Turk Masoor
1.482
19.195**
19.440**
0.269
18.550*
26.088**
31.628**
2.187
7.107
Turk Masoor x Masoor 93
2.871
10.059**
19.440**
0.056
6.304*
11.029**
13.468**
0.740
3.732
ILL 4605 x PL 406
0.050
60.079**
153.520**
71.003**
1.984
6.492*
67.396**
0.979
18.252
PL 406 x ILL 4605
0.456
56.148**
153.520**
61.420**
0.000
8.161**
57.638**
0.579
16.908
** and *: Significant at P < 0.01 and P < 0.05 respectively
216
Appendix IX: Mean square values for nodes on primary branch in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Ps, F1s
Error
Total
vs BCs, F2s Degree of Freedom
2
5
1
ILL 5782 x ILL 2580
1.663
0.978
2.535**
ILL 2580 x ILL 5782
0.294
2.927*
2.535**
ILL 6024 x ILL 8117
3.018
26.694**
LL 8117 x ILL6024
0.820
ILL 6468 x ILL 7556
1
1
1
1
10
17
1.027
0.107
1.125*
0.094
0.219
0.612
9.827*
0.220
0.323
1.730
0.478
1.177
118.637**
0.021
7.482
0.058
7.271
1.694
9.203
27.857**
118.637**
1.491
5.802
2.435
10.920*
1.239
9.018
0.292
20.639**
22.815**
59.042**
2.600*
0.010
18.727**
0.495
6.396
ILL 7556 x ILL 6468
0.072
23.686**
22.815**
64.222**
1.127
7.553*
22.714**
1.13
7.64
ILL 6821 x ILL 7715
1.772
11.535**
29.040**
22.445**
0.050
6.020
0.120
1.401
4.425
ILL 7715 x ILL 6821
0.110
15.340**
29.040**
31.469**
0.570
1.307
14.311**
0.736
4.957
Masoor 93 x Turk Masoor
0.967
25.980**
3.527*
64.222**
2.734
4.919
54.497**
1.065
8.382
Turk Masoor x Masoor 93
0.230
23.010**
3.527*
68.056**
0.920
10.811**
31.734**
0.481
7.077
ILL 4605 x PL 406
0.062
3.371*
6.202*
0.802
0.882
1.608
7.360*
0.843
1.495
PL 406 x ILL 4605
0.061
5.334*
6.202*
0.045
1.084
0.858
18.485**
0.923
2.154
** and *: Significant at P < 0.01 and P < 0.05 respectively
217
Appendix X: Mean square values for fertile nodes on primary branch in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Ps, F1s
Error
Total
vs BCs, F2s Degree of Freedom
2
5
1
1
1
1
1
10
17
ILL 5782 x ILL 2580
0.730
11.363**
27.735**
4.109
4.167
15.383**
5.423
1.365
4.231
ILL 2580 x ILL 5782
0.472
7.941**
27.735**
0.027
0.007
11.584**
0.353
0.527
2.701
ILL 6024 x ILL 8117
2.879
14.353**
39.552**
3.384
18.904**
2.936
6.988*
0.777
5.017
LL 8117 x ILL6024
0.820
15.104**
39.552**
9.687**
8.882**
3.627*
13.773**
0.450
4.804
ILL 6468 x ILL 7556
1.851
9.888**
3.082
23.805**
1.260
0.790
20.501**
1.428
3.966
ILL 7556 x ILL 6468
0.071
13.779**
3.082
36.694**
1.450
2.888
24.781**
1.052
4.680
ILL 6821 x ILL 7715
0.470
4.424**
4.770**
13.607**
0.327
3.415*
0.002
0.622
1.722
ILL 7715 x ILL 6821
0.053
5.610**
4.770**
14.670**
2.344*
2.952*
3.311*
0.412
1.899
Masoor 93 x Turk Masoor
0.350
13.617**
2.667
11.681**
8.167*
5.163*
40.410**
1.037
4.656
Turk Masoor x Masoor 93
0.950
11.929**
2.667
6.845**
2.042
0.429
47.661**
0.580
3.961
ILL 4605 x PL 406
0.361
14.283**
40.042**
9.680**
0.000
0.000
21.692**
0.766
4.694
PL 406 x ILL 4605
0.415
15.795**
40.042**
11.520**
0.150
2.761
24.500**
0.303
4.873
** and *: Significant at P < 0.01 and P < 0.05 respectively
218
Appendix XI: Mean square values for nodes on secondary branch in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Ps, F1s
Error
Total
10
17
vs BCs, F2s Degree of Freedom
2
5
1
1
1
1
1
ILL 5782 x ILL 2580
3.073
1.847
1.707
4.909*
0.050
0.631
1.940
0.821
1.388
ILL 2580 x ILL 5782
0.852
2.573
1.707
1.280
0.060
4.205
5.611
1.568
1.779
ILL 6024 x ILL 8117
1.185
25.875**
104.918**
0.013
14.415**
7.920**
2.108
0.527
8.060
LL 8117 x ILL6024
0.020
28.418**
104.918**
1.022
23.010**
10.595**
2.546
0.917
8.900
ILL 6468 x ILL 7556
0.624
26.574**
11.620**
42.781**
0.844
6.420*
71.202**
0.406
8.128
ILL 7556 x ILL 6468
0.250
28.427**
11.620**
38.574**
3.010
19.034**
69.896**
0.855
8.894
ILL 6821 x ILL 7715
2.402
15.544**
6.720**
52.190**
6.304*
12.317**
0.186
1.059
5.477
ILL 7715 x ILL 6821
0.338
25.697**
6.720**
73.407**
23.010**
17.781**
7.566**
0.615
7.959
Masoor 93 x Turk Masoor
0.720
34.705**
37.500**
64.222**
9.754**
3.117*
58.934**
0.382
10.517
Turk Masoor x Masoor 93
0.044
29.278**
37.500**
66.125**
3.920*
0.066
38.779**
0.410
8.858
ILL 4605 x PL 406
0.275
12.142**
32.202**
10.889**
0.304
0.938
16.378**
0.469
3.879
PL 406 x ILL 4605
0.883
11.959**
32.202**
10.276**
2.344*
1.350
13.624**
0.329
3.815
** and *: Significant at P < 0.01 and P < 0.05 respectively
219
Appendix XII: Mean square values for fertile nodes on secondary branch in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Ps, F1s
Error
Total
vs BCs, F2s Degree of Freedom
2
5
1
1
1
1
1
10
17
ILL 5782 x ILL 2580
0.024
5.535**
7.707**
1.869
6.615**
8.215**
3.268**
0.318
1.818
ILL 2580 x ILL 5782
0.060
6.463**
7.707**
4.302*
0.920
19.282**
0.103
0.568
2.242
ILL 6024 x ILL 8117
1.882
15.655**
44.390**
0.026
15.042**
18.524**
0.294
0.335
5.023
LL 8117 x ILL6024
0.323
17.594**
44.390**
1.135
13.802**
27.281**
1.361
0.363
5.426
ILL 6468 x ILL 7556
0.782
17.824**
4.167*
45.125**
0.770
0.016
39.043**
0.727
5.762
ILL 7556 x ILL 6468
0.624
18.715**
4.167*
38.136**
0.540
2.449
48.282**
0.775
6.034
ILL 6821 x ILL 7715
0.336
14.434**
3.920*
59.224**
1.127
6.504**
1.394
0.568
4.619
ILL 7715 x ILL 6821
0.316
14.632**
3.920*
48.184**
1.654
16.150**
3.251*
0.392
4.571
Masoor 93 x Turk Masoor
0.364
17.873**
22.427**
14.222**
0.050
6.907**
45.761**
0.149
5.388
Turk Masoor x Masoor 93
0.136
16.518**
22.427**
14.045**
0.135
2.832
43.152**
0.117
4.943
ILL 4605 x PL 406
0.037
11.461**
17.510**
14.490**
0.050
0.000
25.252**
0.571
3.711
PL 406 x ILL 4605
0.050
9.464**
17.510**
15.961**
0.107
0.188
13.555**
0.261
2.943
** and *: Significant at P < 0.01 and P < 0.05 respectively
220
Appendix XIII: Mean square values for pod length in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Error
Total
1
10
17
Ps, F1s vs BCs, F2s
Degree of Freedom
2
5
1
ILL 5782 x ILL 2580
0.001
0.002*
0.010**
ILL 2580 x ILL 5782
0.000
0.003*
ILL 6024 x ILL 8117
0.001
LL 8117 x ILL6024
1
1
1
0.000
0.000
0.000
0.000
0.000
0.001
0.010**
0.000
0.000
0.005*
0.000
0.001
0.001
0.014**
0.036**
0.026**
0.000
0.007
0.001
0.001
0.005
0.002
0.012**
0.036**
0.011*
0.001
0.014**
0.000
0.001
0.004
ILL 6468 x ILL 7556
0.000
0.003**
0.001*
0.002**
0.000
0.009**
0.003**
0.000
0.001
ILL 7556 x ILL 6468
0.001
0.003**
0.001*
0.002**
0.000
0.009**
0.001
0.000
0.001
ILL 6821 x ILL 7715
0.002
0.004**
0.012**
0.004*
0.003*
0.000
0.002
0.000
0.002
ILL 7715 x ILL 6821
0.001
0.006**
0.012**
0.003*
0.003*
0.006*
0.004*
0.001
0.002
Masoor 93 x Turk Masoor
0.003
0.048**
0.238**
0.003*
0.001
0.000
0.000
0.003
0.016
Turk Masoor x Masoor 93
0.000
0.050**
0.238**
0.004*
0.000
0.004**
0.004**
0.000
0.015
ILL 4605 x PL 406
0.000
0.041**
0.146**
0.020**
0.008**
0.001
0.032**
0.000
0.012
PL 406 x ILL 4605
0.001
0.043**
0.146**
0.009**
0.007**
0.001
0.053**
0.000
0.013
** and *: Significant at P < 0.01 and P < 0.05 respectively
221
Appendix XIV: Mean square values for pod breadth in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Ps, F1s
Error
Total
10
17
vs BCs, F2s Degree of Freedom
2
5
1
1
ILL 5782 x ILL 2580
0.000
0.002**
0.003**
0.004**
ILL 2580 x ILL 5782
0.013
0.015**
0.003**
0.003**
ILL 6024 x ILL 8117
0.000
0.010**
0.021**
0.015**
LL 8117 x ILL6024
0.001
0.007**
0.021**
0.008**
ILL 6468 x ILL 7556
0.000
0.001**
0.001
ILL 7556 x ILL 6468
0.000
0.001**
ILL 6821 x ILL 7715
0.000
ILL 7715 x ILL 6821
1
1
1
0.001*
0.003**
0.000
0.001
0.027**
0.008**
0.031**
0.013
0.013
0.002**
0.009**
0.001*
0.000
0.003
0.001*
0.004*
0.002*
0.001
0.003
0.002**
0.001*
0.000
0.001*
0.000
0.000
0.001
0.000
0.000
0.001*
0.003**
0.000
0.000
0.005**
0.007**
0.003**
0.002**
0.000
0.012**
0.000
0.001
0.000
0.005**
0.007**
0.001**
0.007**
0.001*
0.011**
0.000
0.002
Masoor 93 x Turk Masoor
0.000
0.003**
0.015**
0.000
0.001*
0.000
0.000
0.000
0.001
Turk Masoor x Masoor 93
0.000
0.003**
0.015**
0.000
0.000
0.001*
0.000
0.000
0.001
ILL 4605 x PL 406
0.000
0.014**
0.061**
0.002**
0.006**
0.000
0.001*
0.000
0.004
PL 406 x ILL 4605
0.000
0.014**
0.061**
0.000
0.003**
0.000
0.005*
0.000
0.004
** and *: Significant at P < 0.01 and P < 0.05 respectively
222
0.000
Appendix XV: Mean square values for seeds per pod in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Ps, F1s
Error
Total
vs BCs, F2s Degree of Freedom
2
5
1
1
1
1
1
10
17
ILL 5782 x ILL 2580
0.000
0.176**
0.493**
0.254**
0.003
0.037**
0.093**
0.002
0.527
ILL 2580 x ILL 5782
0.001
0.168**
0.493**
0.209**
0.001
0.082**
0.056**
0.001
0.050
ILL 6024 x ILL 8117
0.007
0.035**
0.079**
0.001
0.062**
0.005
0.027*
0.005
0.014
LL 8117 x ILL6024
0.000
0.037**
0.079**
0.000
0.058**
0.046**
0.003
0.003
0.013
ILL 6468 x ILL 7556
0.000
0.162**
0.476**
0.057**
0.260**
0.005
0.011
0.004
0.050
ILL 7556 x ILL 6468
0.004
0.150**
0.476**
0.040**
0.232**
0.003
0.000
0.002
0.046
ILL 6821 x ILL 7715
0.009
0.034**
0.104**
0.019
0.012
0.010
0.025*
0.005
0.014
ILL 7715 x ILL 6821
0.001
0.041**
0.104**
0.101**
0.000
0.000
0.001
0.004
0.015
Masoor 93 x Turk Masoor
0.000
0.218**
0.936**
0.048**
0.019**
0.012**
0.073**
0.001
0.065
Turk Masoor x Masoor 93
0.002
0.226**
0.936**
0.046**
0.037**
0.008**
0.102**
0.001
0.067
ILL 4605 x PL 406
0.003
0.240**
0.558**
0.243**
0.028**
0.012*
0.360**
0.002
0.072
PL 406 x ILL 4605
0.002
0.218**
0.558**
0.182**
0.018**
0.007
0.326**
0.005
0.067
** and *: Significant at P < 0.01 and P < 0.05 respectively
223
Appendix XVI: Mean square values for pods per plant in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Ps, F1s
Error
Total
10
17
vs BCs, F2s Degree of Freedom
2
5
1
ILL 5782 x ILL 2580
97.200
6123.600*
20155.000**
ILL 2580 x ILL 5782
210.600
9288.200*
20155.000**
ILL 6024 x ILL 8117
117.500
5427.900*
10778.900*
LL 8117 x ILL6024
34.000
10433.700*
ILL 6468 x ILL 7556
69.700
ILL 7556 x ILL 6468
1
1
1
11.600
418.300
1763.000*
8269.800**
297.000
1987.200
112.800
272.000
12956.200**
12944.900**
71.600
2798.700
7.200
4947.900**
11400.500**
247.900
1756.100
10778.900*
29.600
6353.800**
34727.500**
278.500*
51.100
3102.800
4674.500*
4612.100*
2823.800**
127.000
14730.100**
1079.700*
191.400
1495.700
163.700
4767.800*
4612.100*
2864.000**
275.400
13112.800**
2974.500**
116.300
1489.900
ILL 6821 x ILL 7715
38.300
17159.600**
708.500**
78183.800**
1103.000*
2791.800**
3011.100**
185.300
5160.500
ILL 7715 x ILL 6821
92.400*
19685.700**
708.500**
87696.700**
2497.000**
3309.700**
4216.400**
22.100
5813.800
Masoor 93 x Turk Masoor
85.700
4172.000*
2648.100*
224.400
219.600
2152.600*
15615.600**
227.800
1371.200
Turk Masoor x Masoor 93
67.100
3776.400*
2648.100*
395.300**
254.200*
2511.800**
13072.900**
35.700
1139.600
ILL 4605 x PL 406
223.300*
8349.000**
18067.600**
7046.800**
62.400
197.500
16370.800**
45.100
2508.400
PL 406 x ILL 4605
30.700
8416.000**
18067.600**
8138.800**
0.000
138.800
15734.800**
48.900
2507.700
** and *: Significant at P < 0.01 and P < 0.05 respectively
224
1
5.100
Appendix XVII: Mean square values for 100 seed weight in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Error
Total
1
10
17
Ps, F1s vs BCs, F2s
Degree of Freedom
2
5
ILL 5782 x ILL 2580
0.010
1.013**
ILL 2580 x ILL 5782
0.016
ILL 6024 x ILL 8117
1
1
1
3.072**
0.497**
1.220**
0.005
0.269**
0.008
0.304
1.063**
3.072**
0.464**
1.196**
0.010
0.571**
0.012
0.321
0.010
0.043*
0.008
0.163**
0.032
0.004
0.005
0.009
0.019
LL 8117 x ILL6024
0.012
0.055*
0.008
0.156*
0.017
0.091*
0.004
0.016
0.027
ILL 6468 x ILL 7556
0.000
0.113**
0.461**
0.001
0.007**
0.009**
0.086**
0.001
0.034
ILL 7556 x ILL 6468
0.003
0.123**
0.461**
0.001
0.008
0.011
0.134**
0.003
0.038
ILL 6821 x ILL 7715
0.002
0.237**
0.851**
0.063**
0.034*
0.196**
0.043**
0.004
0.073
ILL 7715 x ILL 6821
0.001
0.289**
0.851**
0.017*
0.083**
0.444**
0.049**
0.003
0.087
Masoor 93 x Turk Masoor
0.001
0.330**
0.316**
0.541**
0.000
0.286**
0.507**
0.002
0.098
Turk Masoor x Masoor 93
0.002
0.340**
0.316**
0.525**
0.000
0.240**
0.619**
0.002
0.101
ILL 4605 x PL 406
0.001
1.869**
6.477**
1.721**
0.478**
0.000
0.668**
0.002
0.551
PL 406 x ILL 4605
0.005
1.825**
6.477**
1.670**
0.322**
0.034*
0.622**
0.006
0.541
** and *: Significant at P < 0.01 and P < 0.05 respectively
225
1
Appendix XVIII: Mean square values for biomass per plant in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Ps, F1s
Error
Total
17
vs BCs, F2s Degree of Freedom
2
5
1
1
1
1
1
10
ILL 5782 x ILL 2580
1.049
15.404**
22.099**
46.713**
0.792
7.036**
0.381
1.124
5.315
ILL 2580 x ILL 5782
0.038
13.550**
22.099**
21.994**
0.110
11.894**
11.651**
0.488
4.277
ILL 6024 x ILL 8117
0.380
23.850**
6.427*
2.816
39.990**
69.240**
0.770
1.680
8.050
LL 8117 x ILL6024
0.679
78.620**
6.430*
32.510**
26.730**
327.430**
0.004
1.180
23.890
ILL 6468 x ILL 7556
0.949
32.909**
18.166**
0.048
0.988
92.367**
52.976**
2.001
10.968
ILL 7556 x ILL 6468
2.190
37.574**
18.166**
0.051
0.013
89.076**
80.565**
1.170
11.997
ILL 6821 x ILL 7715
1.314
95.695**
3.132*
420.162**
2.013
42.072**
11.095*
1.562
29.219
ILL 7715 x ILL 6821
1.531
85.867**
3.132*
348.348**
4.429**
71.429**
1.997
0.428
25.687
Masoor 93 x Turk Masoor
0.220
25.330**
3.100
10.250**
0.001
111.590**
1.270
8.220
Turk Masoor x Masoor 93
0.171
26.810**
3.100
3.850**
10.550**
107.410**
0.660
8.290
ILL 4605 x PL 406
0.410
52.520**
93.102**
17.360**
0.080
0.400
151.660**
0.380
15.720
PL 406 x ILL 4605
0.272
59.043**
93.102**
31.482**
0.132
0.063
170.435**
0.892
17.922
** and *: Significant at P < 0.01 and P < 0.05 respectively
226
1.710 9.130**
Appendix XIX: Mean square values for harvest index per plant in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Ps, F1s
Error
Total
vs BCs, F2s Degree of Freedom
2
5
1
1
1
1
1
10
17
ILL 5782 x ILL 2580
1.553
39.225**
26.670**
7.973*
21.717**
7.311
132.455**
1.512
12.609
ILL 2580 x ILL 5782
0.244
83.674**
26.670**
0.127
70.178**
104.411**
216.986**
1.426
25.478
ILL 6024 x ILL 8117
5.660
11.940**
48.650**
0.310
2.720
1.890
6.140
1.370
4.980
LL 8117 x ILL6024
0.914
14.599**
48.650**
11.139**
7.400*
0.902
4.905
1.096
5.046
ILL 6468 x ILL 7556
2.690
23.084**
44.988**
26.389**
43.360**
0.674
0.006
1.208
7.816
ILL 7556 x ILL 6468
1.641
33.036**
44.988**
56.937**
61.526**
1.274
0.456
0.938
10.461
ILL 6821 x ILL 7715
0.980
34.952**
17.944**
36.960**
51.820**
0.063
67.970**
1.021
10.996
ILL 7715 x ILL 6821
1.213
37.800**
17.944**
31.835**
24.367**
10.977**
103.877**
0.861
11.767
Masoor 93 x Turk Masoor
0.152
77.670**
77.760**
193.770**
0.180
58.820**
57.860**
1.890
23.980
Turk Masoor x Masoor 93
0.046
89.371**
77.760**
243.320**
2.748
31.440**
91.580**
0.769
26.743
ILL 4605 x PL 406
2.530
90.350**
188.320**
92.190**
0.140
0.340
170.770**
0.720
27.290
PL 406 x ILL 4605
1.383
77.990**
188.320**
95.464**
4.432
9.839*
91.888**
1.293
23.861
** and *: Significant at P < 0.01 and P < 0.05 respectively
227
Appendix XX: Mean square values for seed yield per plant in 12 lentil crosses Parameter
Replication
Generation
P1 vs P2
Ps vs F1s
BC1s vs BC2s
BC’s vs F2s
Ps, F1s
Error
Total
10
17
vs BCs, F2s Degree of Freedom
2
5
1
1
1
1
1
ILL 5782 x ILL 2580
0.351
5.661**
4.242**
0.742
2.687**
0.180
20.454**
0.256
1.857
ILL 2580 x ILL 5782
0.170
10.485**
4.242**
1.299
6.699**
7.154**
33.032**
0.373
3.323
ILL 6024 x ILL 8117
0.033
3.007**
3.278**
0.488
6.214**
5.001**
0.051
0.153
0.978
LL 8117 x ILL6024
0.010
9.982**
3.278**
2.836**
5.387**
38.395**
0.015
0.107
3.000
ILL 6468 x ILL 7556
0.171
4.688**
6.813**
0.842
0.204
9.405**
6.177**
0.231
1.535
ILL 7556 x ILL 6468
0.227
5.803**
6.813**
1.477*
1.097*
10.661**
8.967**
0.200
1.851
ILL 6821 x ILL 7715
0.049
14.599**
0.634*
63.807**
1.340**
3.923**
3.293**
0.127
4.374
ILL 7715 x ILL 6821
0.113
12.289**
0.634*
52.429**
1.368**
4.952**
2.063**
0.075
3.672
Masoor 93 x Turk Masoor
0.018
5.959**
3.390**
6.907**
0.173
0.201
19.123**
0.185
1.864
Turk Masoor x Masoor 93
0.024
6.035**
3.390**
6.969**
0.608*
0.256
18.950**
0.062
1.815
ILL 4605 x PL 406
0.026
5.940**
14.837**
0.011
0.004
0.028
14.850**
0.020
1.763
PL 406 x ILL 4605
0.002
6.121**
14.837**
0.014
0.003
0.001
15.751**
0.031
1.819
** and *: Significant at P < 0.01 and P < 0.05 respectively
228
Appendix XXI: Joint scaling test for morphological characteristics in 12 lentil crosses Cross
ILL 5782 x ILL 2580 ILL 2580 x ILL 5782
ILL 6024 x ILL 8117 ILL 8117 x ILL 6024
ILL 6468 x ILL 7556 ILL 7556 x ILL 6468
ILL 6821 x ILL 7715 ILL 7715 x ILL 6821
Masoor 93 x Turk Masoor Turk Masoor x Masoor 93
ILL 4605 x PL 406 PL 406 x ILL 4605
Estimate
Days to flower
Days to mature
Plant height
Primary branches plant-1
Secondary branches plant-1
Nodes on primary branch
Fertile nodes on primary branch
Nodes on secondary branch
Fertile nodes on secondary branch
m d h m d h
98.30±0.20 0.52±0.21 4.99±0.45 98.08±0.21 0.66±0.21 4.45±0.49
151.38±0.18 –1.79±0.19 2.16±0.32 151.42±0.18 –1.86±0.19 2.01±0.30
37.65±0.33 1.11±0.32 –1.08±0.68 36.32±0.32 1.81±0.32 –0.84±0.58
2.77±0.05 0.16±0.05 0.04±0.09 2.74±0.05 0.20±0.05 –0.10±0.09
14.45±0.30 3.71±0.29 3.05±0.64 13.60±0.29 3.17±0.28 1.88±0.58
21.94±0.28* 0.57±0.27 –0.72±0.52 21.74±0.26* 0.40±0.27 –2.38±0.50
11.54±0.23 2.15±0.23 1.45±0.45 11.01±0.22 1.54±0.23 –0.19±0.40
18.15±0.25* 1.30±0.25 –0.62±0.48 18.66±0.25* 0.43±0.25 –2.01±0.51
m d h m d h
97.96±0.17 –0.48±0.18 5.06±0.33 97.82±0.177 –0.56±0.18 4.83±0.33
147.09±0.12 3.87±0.12 –5.43±0.30 147.03±0.12 3.87±0.12 –5.86±0.31
36.39±0.26 7.53±0.26 –6.17±0.78 35.37±0.26 6.18±0.26 –7.71±0.54
2.57±0.04 0.17±0.04 –0.16±0.07 2.59±0.04 –0.07±0.04 –0.26±0.08
15.46±0.26 3.27±0.27 –2.32±0.44 13.98±0.25 1.55±0.26 –1.62±0.44
20.96±0.25 4.13±0.26 –0.59±0.48 21.19±0.25 3.50±0.26 –1.51±0.48
12.41±0.21 2.75±0.21 –2.09±0.42 12.17±0.20 1.50±0.21 –2.38±0.35
17.26±0.20 3.94±0.21 –0.43±0.37 17.05±0.20 3.09±0.21 –0.95±0.37
9.06±0.18 2.85±0.17 –0.38±0.36 8.31±0.16 1.17±0.17 –0.31±0.26
m d h m d h
98.96±0.22 2.59±0.20 0.34±0.41 98.67±0.23 2.27±0.22 –0.22±0.44
140.32±0.11 1.02±0.11 –5.76±0.21 140.23±0.12 0.92±0.11 –5.56±0.27
30.45±0.25 4.87±0.24 –6.53±0.47 30.84±0.26 4.18±0.25 –5.66±0.50
2.51±0.05 –0.09±0.05 –0.08±0.10 2.54±0.05 –0.11±0.05 –0.02±0.10
14.82±0.25 1.25±0.25 – 0.63±0.53 14.66±0.25 0.78±0.25 –0.84±0.53
21.29±0.18 1.96±0.18 –6.44±0.40 21.38±0.18 1.63±0.19 –6.20±0.37
13.12±0.21 –1.48±0.20 –9.35±0.44 13.12±0.20 0.96±0.20 –5.03±0.40
16.95±0.17 1.08±0.16 –6.97±0.35 17.18±0.17 0.71±0.17 –6.39±0.35
9.66±0.20 0.79±0.18 –6.00±0.39 8.96±0.19 0.09±0.17 –4.85±0.35
m d h m d h
79.82±0.33 13.70±0.34 17.26±0.43 79.56±0.34 13.88±0.34 17.19±0.43
133.22±0.12 6.50±0.12 5.17±0.26 133.21±0.12 6.50±0.12 5.33±0.26
25.66±0.28 –3.09±0.29 12.78±0.60 25.60±0.28 3.40±0.29 11.43±0.61
2.25±0.04 0.33±0.04 0.30±0.09 2.29±0.04 0.22±0.05 0.41±0.09
8.86±0.20 0.72±0.21 5.16±0.42 6.72±1.00 0.65±0.20 4.24±0.37
18.61±0.21 –1.93±0.21 2.75±0.46 18.02±0.21 –1.94±0.21 1.48±0.44
10.14±0.20 –0.65±0.20 2.03±0.42 9.85±0.20 –0.79±0.20 1.62±0.39
12.69±0.20 1.14±0.20 4.13±0.42 12.20±0.19 0.64±0.20 4.04±0.40
5.89±0.15 0.78±0.16 4.26±0.31 5.57±0.15 0.53±0.16 2.96±0.31
m d h m d h
91.13±0.23 –5.76±0.23 7.55±0.44 91.42±0.23 –5.40±0.22 6.73±0.42
143.99±0.13 –8.20±0.13 –10.98±0.25 144.15±0.18 –8.37±0.19 –11.13±0.30
34.24±0.26 –1.63±0.27 –9.34±0.51 34.65±0.26 –1.43±0.27 –6.92±0.51
2.66±0.04 0.00±0.04 –0.34±0.08 2.73±0.04* 0.02±0.04 –0.19±0.08
13.48±0.20 –1.40±0.21 –0.30±0.35 13.40±0.20 –0.95±0.21 0.20±0.35
20.86±0.20 – 0.35±0.20 – 6.33±0.36 21.08±0.19 –0.29±0.20 –5.85±0.34
8.49±0.20 2.17±0.20 –5.75±0.36 11.39±0.17 0.30±0.17 –2.12±0.29
17.76±0.17 –1.99±0.17 –6.38±0.31 17.91±0.17 –1.46±0.18 –5.95±0.32
7.90±0.14 – 0.18±0.14 – 1.41±0.23 7.90±0.15 –0.49±0.15 –1.35±0.23
m d h m d h
88.89±0.21 8.04±0.21 23.35±0.71 90.06±0.31 –5.44±0.27 11.67±0.54
134.54±0.11 9.34±0.11 14.37±0.33 134.54±0.11 9.33±0.11 13.96±0.35
26.91±0.22 2.92±0.23 3.21±0.41 27.21±0.23 2.80±0.23 4.64±0.41
2.40±0.06 0.03±0.05 0.23±0.10 2.34±0.05 0.04±0.05 –0.05±0.09
8.74±0.29 3.31±0.29 2.79±0.58 8.72±0.29 3.40±0.29 2.40±0.57
18.29±0.19 0.88±0.19 –0.28±0.41 18.05±0.19 1.06±0.19 –2.21±0.42
8.22±0.17 2.42±0.18 1.68±0.29 7.69±0.16 2.41±0.18 1.57±0.29
13.37±0.17 2.21±0.17 0.70±0.40 13.43±0.17 2.08±0.17 0.50±0.40
5.00±0.15 1.05±0.15 1.16±0.29 5.14±0.15 1.09±0.15 1.25±0.30
*: Non-significant χ2 test
m: Mean effect, d: Additive gene effect,
h: Dominance gene effect
229
8.14±0.17 1.41±0.18 1.01±0.31 7.59±0.17 0.66±0.17 1.25±0.33
Appendix XXII: Joint scaling test for yield and yield related characteristics in 12 lentil crosses Cross ILL 5782 x ILL 2580 ILL 2580 x ILL 5782
ILL 6024 x ILL 8117 ILL 8117 x ILL 6024
ILL 6468 x ILL 7556 ILL 7556 x ILL 6468
ILL 6821 x ILL 7715 ILL 7715 x ILL 6821
Masoor 93 x Turk Masoor Turk Masoor x Masoor 93
ILL 4605 x PL 406 PL 406 x ILL 4605
Estimate
Pod length
Pod breadth
Seed pod-1
Pods plant-1
100 seed weight
Biomass
Harvest index
Seed yield plant-1
m d h m d h
1.04±0.01 –0.02±0.01 –0.01±0.01 1.05±0.01 –0.04±0.01 –0.01±0.01
0.54±0.01 –0.01±0.01 0.05±0.01 0.53±0.01 –0.02±0.01 0.06±0.01
1.56±0.02 0.23±0.02 –0.39±0.03 1.59±0.02 0.22±0.02 –0.34±0.03
196.24±3.23 51.27±3.27 –27.18±6.53 184.41±3.16 48.52±3.23 –35.49±5.96
2.59±0.01 –0.69±0.01 0.39±0.03 2.43±0.01 –0.51±0.01 0.29±0.03
16.87±0.36 1.19±0.36 –2.43±0.75 14.61±0.34 3.12±0.34 –5.76±0.68
35.20±0.33 1.72±0.33 –4.73±0.61 34.81±0.32 1.77±0.33 –3.50±0.55
5.54±0.20 0.70±0.20 –1.01±0.32 4.06±0.18 2.46±0.17 0.03±0.32
m d h m d h
1.02±0.01 –0.06±0.01 0.07±0.01 1.03±0.01 –0.07±0.01 0.04±0.01
0.56±0.00 –0.05±0.00 0.07±0.01 0.58±0.01 –0.05±0.01 0.08±0.01
1.35±0.03 –0.01±0.02 –0.05±0.04 1.27±0.02 –0.15±0.02 –0.00±0.04
173.96±2.89 39.74±2.93 –1.81±5.57 153.37±2.76 32.53±2.88 –22.38±5.25
2.62±0.01 0.02±0.01 0.28±0.03 2.61±0.01 0.04±0.01 0.22±0.03
11.81±0.29 1.45±0.30 0.78±0.53 9.34±0.28 –1.18±0.28 1.06±0.55
35.64±0.32 2.69±0.32 –0.73±0.60 35.64±0.32 2.20±0.32 –3.04±0.68
4.22±0.11 0.83±0.11 0.36±0.21 3.49±0.10 0.30±0.10 –0.22±0.21
m d h m d h
1.04±0.01 0.01±0.01 0.03±0.01 1.03±0.01 0.01±0.01 0.02±0.01
0.55±0.00 –0.01±0.00 –0.03±0.01 0.53±0.01 –0.01±0.01 –0.02±0.01
1.58±0.01 0.30±0.01 0.13±0.03 1.64±0.01 0.20±0.01 0.07±0.03
195.40±3.37 15.81±3.41 –35.91±5.67 191.90±3.31 43.49±3.33 –41.68±3.84
2.08±0.01 –0.26±0.01 –0.01±0.02 2.06±0.01 –0.24±0.01 –0.08±0.02
13.47±0.35 0.76±0.35 –2.02±0.66 11.51±0.31 –0.06±0.31 1.30±0.51
36.54±0.37 3.11±0.37 –3.40±0.68 36.66±0.37 1.73±0.38 –5.15±0.33
5.07±0.12 0.71±0.12 – 0.95±0.22 4.49±0.11 0.06±0.11 –0.50±0.19
m d h m d h
1.15±0.00 –0.05±0.00 –0.02±0.01 1.15±0.01 –0.04±0.01 –0.01±0.01
0.55±0.00 –0.04±0.00 0.06±0.01 0.54±0.01 –0.03±0.01 0.05±0.01
1.61±0.02 0.14±0.02 –0.15±0.03 1.65±0.02* –0.12±0.02 –0.21±0.03
54.74±1.43 12.29±1.44 144.33±4.69 52.72±1.42 11.68±1.44 148.54±4.08
2.62±0.02 –0.30±0.02 0.22±0.03 2.57±0.02 –0.22±0.02 0.19±0.03
5.11±0.14 0.89±0.15 8.89±0.44 5.12±0.14 0.82±0.15 8.42±0.42
31.73±0.40 0.42±0.42 0.63±0.63 31.72±0.40 1.41±0.42 3.96±0.62
1.73±0.05 0.38±0.05 3.09±0.18 1.71±0.06 0.35±0.06 3.14±0.16
m d h m d h
1.13±0.01 0.19±0.01 –0.03±0.01 1.12±0.01 0.18±0.01 –0.08±0.01
0.55±0.00 0.04±0.00 0.01±0.01 0.55±0.01 0.05±0.01 –0.01±0.01
1.54±0.01 0.37±0.01 –0.22±0.03 1.51±0.01 0.39±0.01 –0.30±0.03
152.41±2.53 –20.55±2.55 –19.87±5.01 147.81±2.47 –13.58±2.50 –11.37±4.72
2.59±0.01 0.17±0.01 –0.53±0.02 2.58±0.01 0.18±0.01 –0.51±0.02
10.01±0.22 –0.17±0.23 –1.68±0.33 10.59±0.24 0.83±0.23 –4.92±0.46
39.06±0.6 3.27±0.36 –11.48±0.78 38.98±0.35 3.23±0.35 –12.62±0.72
3.93±0.09 0.17±0.09 –1.95±0.15 3.91±0.09 0.55±0.09 –2.03±0.15
m d h m d h
1.15±0.00 –0.15±0.00 –0.14±0.01 1.14±0.01 –0.14±0.01 –0.14±0.01
0.60±0.00 –0.10±0.00 –0.04±0.01 0.60±0.01 –0.10±0.01 –0.05±0.01
1.56±0.01 0.33±0.01 –0.50±0.03 1.59±0.02 0.29±0.02 –0.39±0.03
73.98±1.85 43.81±1.87 –4.39±4.06 69.25±1.82 38.69±1.85 17.52±3.77
2.88±0.01 –1.02±0.01 –1.05±0.03 2.85±0.01 –0.99±0.01 –1.02±0.03
6.73±0.17 3.02±0.16 –4.47±0.36 6.47±0.17 2.59±0.16 –3.50±0.37
27.82±0.32 5.52±0.34 –7.01±0.52 28.79±0.34 5.45±0.34 –7.30±0.58
1.96±0.05 1.21±0.05 –1.66±0.10 1.79±0.05 0.99±0.05 –1.28±0.10
*: Non-significant χ2 test
m: Mean effect, d: Additive gene effect,
h: Dominance gene effect
230
Appendix XXIII: Mean values and standard deviations for different plant traits in cross ILL 5782 x ILL 2580 Parameter
ILL 5782
ILL 2580
F1 generation
F2 generation
Days to flower
97.25
±
0.65
98.63
±
0.73
101.75
±
1.52
102.05
±
0.94
Days to mature
152.28
±
0.46
147.68
±
0.48
151.78
±
0.7
155.45
±
2.56
Plant height (cm)
36.67
±
0.08
39.92
±
1.03
39.42
±
0.38
35.98
±
0.93
Primary branches
2.65
±
0.18
3.08
±
0.10
3.08
±
0.06
2.67
±
0.04
Secondary branches
10.67
±
0.75
19.71
±
0.4
19.68
±
2.43
14.33
±
1.88
Nodes on primary branch
21.30
±
0.74
22.6
±
0.48
21.23
±
0.81
22.07
±
0.55
Fertile nodes on primary branch
9.38
±
0.60
13.68
±
0.37
12.97
±
0.2
11.26
±
1.11
Nodes on secondary branch
18.65
±
0.22
19.72
±
1.69
17.62
±
0.76
17.63
±
1.59
Fertile nodes on secondary branch
6.82
±
0.38
9.08
±
0.48
8.92
±
0.31
7.77
±
0.84
Pod length (cm)
1.08
±
0.02
1.00
±
0.00
1.03
±
0.01
1.03
±
0.05
Pod breadth (cm)
0.54
±
0.01
0.50
±
0.00
0.57
±
0.02
0.54
±
0.03
Seeds pod-1
1.30
±
0.02
1.87
±
0.03
1.23
±
0.05
1.41
±
0.03
Pods plant-1
151.4
±
3.49
267.32
±
11.36
205.99
±
18.23
145.89
±
26.25
100 seed weight (g)
3.35
±
0.07
1.92
±
0.02
3.14
±
0.01
2.59
±
0.18
Biomass plant-1 (g)
16.30
±
0.63
20.52
±
0.98
20.4
±
0.93
12.37
±
2.04
Harvest index plant-1
33.11
±
0.51
36.95
±
0.62
30.12
±
0.82
34.38
±
1.57
Yield plant-1 (g)
5.88
±
0.82
7.56
±
0.53
6.06
±
0.47
4.18
±
0.51
Appendix XXIV: Mean values and standard deviations for different plant traits in cross ILL 2580 x ILL 5782 Parameter
ILL 2580
ILL 5782
F1 generation
F2 generation
Days to flower
101.73
±
1.19
97.25
±
0.65
101.73
±
1.19
99.4
±
0.46
Days to mature
151.9
±
0.74
152.28
±
0.46
151.9
±
0.74
155.48
±
2.61
Plant height (cm)
39.0
±
0.05
36.67
±
0.08
39.00
±
0.05
32.63
±
1.14
Primary branches
3.05
±
0.05
2.65
±
0.18
3.05
±
0.05
2.55
±
0.12
Secondary branches
18.88
±
0.35
10.67
±
0.75
18.88
±
0.35
10.14
±
1.67
Nodes on primary branch
19.73
±
0.68
21.3
±
0.74
19.73
±
0.68
20.32
±
1.01
Fertile nodes on primary branch
11.42
±
0.52
9.38
±
0.6
11.42
±
0.52
9.61
±
1.01
Nodes on secondary branch
18.38
±
2.14
18.65
±
0.22
18.38
±
2.14
16.83
±
0.98
Fertile nodes on secondary branch
9.42
±
1.29
6.82
±
0.38
9.42
±
1.29
6.52
±
0.86
Pod length (cm)
1.03
±
0.01
1.08
±
0.02
1.03
±
0.01
1.08
±
0.05
Pod breadth (cm)
0.56
±
0.02
0.54
±
0.01
0.56
±
0.02
0.57
±
0.04
Seeds pod-1
1.26
±
0.04
1.30
±
0.02
1.26
±
0.04
1.50
±
0.03
Pods plant-1
151.40
±
3.49
201.58
±
6.86
201.58
±
6.86
99.01
±
18.54
100 seed weight (g)
3.35
±
0.07
3.12
±
0.04
3.12
±
0.04
2.39
±
0.04
Biomass plant-1 (g)
16.30
±
0.63
18.66
±
0.50
18.66
±
0.50
6.69
±
1.18
Harvest index plant-1
33.11
±
0.51
31.71
±
0.70
31.71
±
0.70
33.94
±
0.90
Yield plant-1 (g)
5.88
±
0.82
5.91
±
0.30
5.91
±
0.30
2.48
±
0.38
231
Appendix XXV: Mean values and standard deviations for different plant traits in cross ILL 6024 x ILL 8117 Parameter
ILL 6024
ILL 8117
F1 generation
F2 generation
Days to flower
98.05
±
0.30
97.32
±
0.53
102.25
±
0.70
100.81
±
0.59
Days to mature
142.42
±
0.18
151.15
±
0.15
142.27
±
0.93
146.74
±
3.09
Plant height (cm)
29.02
±
0.19
44.55
±
0.78
31.07
±
0.6
32.26
±
1.55
Primary branches
2.58
±
0.06
2.69
±
0.01
2.53
±
0.25
2.38
±
0.13
Secondary branches
12.77
±
0.57
19.43
±
2.27
14.22
±
0.41
12.33
±
1.01
Nodes on primary branch
17.03
±
1.15
25.93
±
1.11
21.58
±
2.64
20.13
±
1.14
Fertile nodes on primary branch
10.28
±
0.85
15.42
±
0.98
11.55
±
1.43
10.36
±
0.73
Nodes on secondary branch
13.6
±
0.35
21.96
±
0.63
17.7
±
0.97
15.74
±
1.17
Fertile nodes on secondary branch
6.50
±
1.02
11.94
±
0.81
9.33
±
1.11
6.97
±
0.34
Pod length (cm)
1.12
±
0.02
0.96
±
0.01
1.16
±
0.07
1.02
±
0.02
Pod breadth (cm)
0.64
±
0.02
0.52
±
0.01
0.66
±
0.01
0.57
±
0.02
Seeds pod-1
1.40
±
0.09
1.17
±
0.02
1.26
±
0.03
1.32
±
0.12
Pods plant-1
15.61
134.62
±
7.51
219.39
±
11.83
178.9
±
23.5
128.37
±
100 seed weight (g)
2.58
±
0.06
2.66
±
0.03
2.91
±
0.12
2.79
±
0.15
Biomass plant-1 (g)
10.94
±
0.55
13.01
±
1.20
13.16
±
1.58
8.86
±
1.18
Harvest index plant-1
33.06
±
1.36
38.76
±
1.28
35.55
±
2.04
35.26
±
0.91
3.51
±
0.26
4.99
±
0.36
4.74
±
0.35
3.47
±
0.48
-1
Seed yield plant (g)
Appendix XXVI: Mean values and standard deviations for different plant traits in cross ILL 8117 x ILL 6024 Parameter
ILL 8117 ± 0.53
F1 generation
F2 generation
98.05
ILL 6024 ±
0.30
102.18
±
0.78
100.68
±
0.35
0.15
142.42
±
0.18
142.6
±
1.01
146.47
±
2.30
±
0.78
29.02
±
0.19
31.23
±
0.9
29.68
±
0.26
±
0.01
2.58
±
0.06
2.50
±
0.05
2.33
±
0.04
19.43
±
2.27
12.77
±
0.57
14.88
±
1.63
10.28
±
0.16
Nodes on primary branch
25.93
±
1.11
17.03
±
1.15
20.62
±
1.35
20.37
±
0.47
Fertile nodes on primary branch
15.42
±
0.98
10.28
±
0.85
10.65
±
0.53
9.47
±
0.68
Days to flower
97.32
Days to mature
151.15
±
Plant height (cm)
44.55
Primary branches
2.69
Secondary branches
Nodes on secondary branch
21.96
±
0.63
13.6
±
0.35
17.07
±
0.71
15.26
±
0.59
Fertile nodes on secondary branch
11.94
±
0.81
6.50
±
1.02
8.47
±
0.18
5.96
±
0.38
Pod length (cm)
0.96
±
0.01
1.12
±
0.02
1.12
±
0.07
1.01
±
0.01
Pod breadth (cm)
0.52
±
0.01
0.64
±
0.02
0.64
±
0.00
0.59
±
0.05
Seeds pod-1
1.17
±
0.02
1.40
±
0.09
1.28
±
0.02
1.21
±
0.04 2.24
-1
219.39
±
11.83
134.62
±
7.51
180.85
±
7.16
81.93
±
100 seed weight (g)
2.66
±
0.03
2.58
±
0.06
2.91
±
0.11
2.55
±
0.25
Biomass plant-1 (g)
13.01
±
1.20
10.94
±
0.55
16.01
±
1.89
4.78
±
0.26
Harvest index plant-1
38.76
±
1.28
33.06
±
1.36
33.55
±
1.04
33.63
±
1.27
Seed yield plant-1 (g)
4.99
±
0.36
3.51
±
0.26
5.44
±
0.42
1.67
±
0.10
Pods plant
232
Appendix XXVII: Mean values and standard deviations for different plant traits in cross ILL 6468 x ILL 7556 Parameter
ILL 6468
ILL 7556
F1 generation
F2 generation
Days to flower
99.03
±
0.13
100.33
±
0.53
99.88
±
0.63
98.75
±
1.24
Days to mature
139.52
±
0.59
141.1
±
0.35
134.48
±
0.40
138.33
±
2.64
Plant height (cm)
25.88
±
0.26
36.90
±
0.65
26.57
±
0.13
26.29
±
2.13
Primary branches
2.85
±
0.22
2.47
±
0.12
2.88
±
0.13
2.25
±
0.11
Secondary branches
14.03
±
0.60
16.25
±
1.31
15.47
±
0.78
12.97
±
1.65
Nodes on primary branch
19.57
±
0.89
23.47
±
0.72
16.08
±
0.89
17.9
±
0.93
Fertile nodes on primary branch
12.92
±
0.86
14.35
±
0.56
10.18
±
2.68
9.74
±
0.67
Nodes on secondary branch
16.72
±
0.58
19.50
±
0.70
13.48
±
0.46
14.02
±
1.59
Fertile nodes on secondary branch
9.75
±
0.96
11.42
±
1.03
5.83
±
0.51
5.89
±
0.59
Pod length (cm)
1.02
±
0.01
1.05
±
0.02
1.06
±
0.01
1.02
±
0.03
Pod breadth (cm)
0.57
±
0.01
0.55
±
0.03
0.52
±
0.01
0.52
±
0.02
Seeds pod-1
1.32
±
0.06
1.88
±
0.04
1.77
±
0.07
1.52
±
0.10
Pods plant-1
180.65
±
9.90
236.10
±
15.84
170.8
±
7.87
122.72
±
22.75
100 seed weight (g)
2.38
±
0.03
1.83
±
0.02
2.13
±
0.02
2.01
±
0.04
Biomass plant-1 (g)
13.98
±
1.46
17.46
±
1.45
15.57
±
1.13
7.72
±
2.18
Harvest index plant-1
33.67
±
1.58
39.14
±
0.68
32.77
±
1.00
33.72
±
4.88
4.64
±
0.43
6.77
±
0.63
5.06
±
0.52
2.75
±
0.64
-1
Seed yield plant (g)
Appendix XXVIII: Mean values and standard deviations for different plant traits in cross ILL 7556 x ILL 6468 Parameter
ILL 7556
ILL 6468
F1 generation
F2 generation
Days to flower
100.33
±
0.53
99.03
±
0.13
99.7
±
2.76
97.99
±
5.90
Days to mature
141.1
±
0.35
139.52
±
0.59
135.1
±
2.43
138.15
±
2.67
Plant height (cm)
36.9
±
0.65
25.88
±
0.26
27.52
±
3.53
29.48
±
37.2
Primary branches
2.47
±
0.12
2.85
±
0.22
2.77
±
0.70
2.44
±
0.57
Secondary branches
16.25
±
1.31
14.03
±
0.6
15.83
±
4.25
13.10
±
5.42
Nodes on primary branch
23.47
±
0.72
19.57
±
0.89
15.85
±
2.79
18.68
±
3.40
Fertile nodes on primary branch
14.35
±
0.56
12.92
±
0.86
9.35
±
2.79
10.66
±
3.51
Nodes on secondary branch
19.5
±
0.7
16.72
±
0.58
13.72
±
2.66
14.76
±
3.13
Fertile nodes on secondary branch
11.42
±
1.03
9.75
±
0.96
6.22
±
2.10
6.59
±
2.84
Pod length (cm)
1.05
±
0.02
1.02
±
0.01
1.07
±
0.09
1.01
±
0.09
Pod breadth (cm)
0.55
±
0.03
0.57
±
0.01
0.56
±
0.05
0.51
±
0.04
Seeds pod-1
1.88
±
0.04
1.32
±
0.06
1.74
±
0.20
1.67
±
0.34
Pods plant-1
67.59
236.10
±
15.84
180.65
±
9.9
170.5
±
36.37
116.07
±
100 seed weight (g)
1.83
±
0.02
2.38
±
0.03
2.08
±
0.14
1.88
±
0.33
Biomass plant-1 (g)
17.46
±
1.45
13.98
±
1.46
15.56
±
2.56
6.99
±
4.52
Harvest index plant-1
39.14
±
0.68
33.67
±
1.58
31.07
±
5.14
34.41
±
9.46
Seed yield plant-1 (g)
6.77
±
0.63
4.64
±
0.43
4.85
±
1.03
2.47
±
1.82
233
Appendix XXIX: Mean values and standard deviations for different plant traits in cross ILL 6821 x ILL 7715 Parameter
ILL 6821
ILL 7715
F1 generation
F2 generation
Days to flower
65.53
±
4.81
93.27
±
2.24
95.43
±
10.54
90.69
±
18.07
Days to mature
126.75
±
0.93
139.93
±
1.67
136.77
±
15.70
132.47
±
10.97
Plant height (cm)
29.77
±
3.32
22.83
±
3.20
40.38
±
6.91
30.17
±
6.36
Primary branches
2.03
±
0.55
2.63
±
0.49
2.79
±
0.64
2.10
±
0.90
Secondary branches
8.78
±
2.64
9.87
±
1.98
15.36
±
3.42
9.29
±
5.25
Nodes on primary branch
21.02
±
2.66
16.62
±
2.24
21.87
±
4.44
18.94
±
4.78
Fertile nodes on primary branch
11.28
±
2.66
9.50
±
2.00
12.87
±
3.52
10.41
±
3.67
Nodes on secondary branch
11.93
±
2.26
14.05
±
2.27
17.90
±
3.64
13.24
±
4.76
Fertile nodes on secondary branch
5.60
±
1.73
7.22
±
1.89
11.77
±
2.37
6.46
±
3.34
Pod length (cm)
1.19
±
0.07
1.10
±
0.03
1.21
±
0.89
1.15
±
0.15
Pod breadth (cm)
0.57
±
0.04
0.51
±
0.02
0.71
±
1.08
0.60
±
0.08
Seeds pod-1
1.50
±
0.22
1.77
±
0.14
1.68
±
1.11
1.48
±
0.28
Pods plant-1
47.90
±
18.36
69.63
±
13.04
252.46
±
52.74
73.90
±
68.31
100 seed weight (g)
3.08
±
0.32
2.33
±
0.13
3.03
±
1.18
2.46
±
0.50
Biomass plant-1 (g)
4.81
±
1.90
6.26
±
1.28
19.93
±
4.34
5.74
±
5.25
Harvest index plant-1
31.62
±
5.11
35.08
±
4.58
37.27
±
4.52
30.77
±
9.52
1.54
±
0.71
2.19
±
0.50
7.60
±
2.06
1.96
±
2.09
-1
Seed yield plant (g)
Appendix XXX: Mean values and standard deviations for different plant traits in cross ILL 7715 x ILL 6821 Parameter
ILL 7715
ILL 6821
F1 generation
F2 generation
Days to flower
93.27
±
2.24
65.53
±
4.81
96.58
±
2.05
89.03
±
17.53
Days to mature
139.93
±
1.67
126.75
±
0.93
138.93
±
1.76
131.95
±
10.91
Plant height (cm)
22.83
±
3.20
29.77
±
3.32
39.97
±
4.73
28.38
±
6.91
Primary branches
2.63
±
0.49
2.03
±
0.55
2.90
±
0.71
2.18
±
0.89
Secondary branches
9.87
±
1.98
8.78
±
2.64
14.30
±
2.41
8.69
±
4.37
Nodes on primary branch
16.62
±
2.24
21.02
±
2.66
22.78
±
3.59
17.82
±
3.13
Fertile nodes on primary branch
9.50
±
2.00
11.28
±
2.66
13.10
±
2.86
9.63
±
3.45
Nodes on secondary branch
14.05
±
2.27
11.93
±
2.26
19.05
±
3.02
11.73
±
4.15
Fertile nodes on secondary branch
7.22
±
1.89
5.60
±
1.73
11.32
±
2.36
5.30
±
2.58
Pod length (cm)
1.10
±
0.03
1.19
±
0.07
1.11
±
0.11
1.13
±
0.11
Pod breadth (cm)
0.51
±
0.02
0.57
±
0.04
0.57
±
0.05
0.58
±
0.06
Seeds pod-1
1.77
±
0.14
1.50
±
0.22
1.41
±
0.25
1.55
±
0.31
Pods plant-1
69.63
±
13.04
47.90
±
18.36
268.17
±
36.18
70.84
±
56.72
100 seed weight (g)
2.33
±
0.13
3.08
±
0.32
2.80
±
0.14
2.32
±
0.45
Biomass plant-1 (g)
6.26
±
1.28
4.81
±
1.90
18.73
±
3.91
5.28
±
4.94
Harvest index plant-1
35.08
±
4.58
31.62
±
5.11
37.34
±
3.41
31.43
±
9.28
Seed yield plant-1 (g)
2.19
±
0.50
1.54
±
0.71
6.99
±
1.56
1.85
±
1.98
234
Appendix XXXI: Mean values and standard deviations for different plant traits in cross Masoor 93 x Tuk Masoor Parameter
Masoor 93
Turk Masoor
F1 generation
F2 generation
Days to flower
97.02
±
1.48
79.12
±
3.98
95.67
±
3.00
96.89
±
6.46
Days to mature
151.15
±
1.72
136.62
±
1.17
133.85
±
1.82
132.78
±
5.53
Plant height (cm)
37.20
±
3.30
33.62
±
2.79
28.10
±
3.56
26.10
±
6.53
Primary branches
2.72
±
0.45
2.73
±
0.45
2.53
±
0.57
2.34
±
0.58
Secondary branches
16.10
±
3.22
12.50
±
1.43
14.67
±
2.15
9.36
±
5.80
Nodes on primary branch
22.37
±
3.19
20.83
±
1.40
15.93
±
2.25
17.28
±
3.80
Fertile nodes on primary branch
13.60
±
2.57
12.27
±
1.59
10.52
±
1.95
8.06
±
3.71
Nodes on secondary branch
21.52
±
2.70
16.52
±
1.52
13.35
±
1.87
12.68
±
3.38
Fertile nodes on secondary branch
12.33
±
2.71
8.47
±
1.43
7.73
±
1.15
5.08
±
2.76
Pod length (cm)
0.92
±
0.07
1.32
±
0.05
1.08
±
0.09
1.10
±
0.11
Pod breadth (cm)
0.50
±
0.00
0.60
±
0.00
0.55
±
0.05
0.56
±
0.07
Seeds pod-1
1.14
±
0.19
1.93
±
0.10
1.38
±
0.25
1.40
±
0.63
Pods plant-1
71.78
188.55
±
31.37
146.53
±
27.34
178.13
±
35.85
90.29
±
100 seed weight (g)
2.44
±
0.14
2.90
±
0.16
2.15
±
0.12
2.41
±
0.45
Biomass plant-1 (g)
11.53
±
2.54
12.97
±
2.90
9.99
±
1.69
6.50
±
4.78
Harvest index plant-1
35.63
±
5.07
42.83
±
2.99
29.39
±
6.05
35.98
±
14.23
4.06
±
0.88
5.56
±
1.32
2.95
±
0.89
2.34
±
1.90
-1
Seed yield plant (g)
Appendix XXXII: Mean values and standard deviations for different plant traits in cross Turk Masoor x Masoor 93 Parameter
Turk Masoor
Masoor 93
F1 generation
F2 generation
Days to flower
79.12
±
3.98
97.02
±
1.48
95.48
±
2.78
96.34
±
5.47
Days to mature
136.62
±
1.17
151.15
±
1.72
133.55
±
1.87
133.18
±
5.56
Plant height (cm)
33.62
±
2.79
37.20
±
3.30
27.77
±
3.61
29.10
±
5.49
Primary branches
2.73
±
0.45
2.72
±
0.45
2.53
±
0.57
2.61
±
1.33
Secondary branches
12.50
±
1.43
16.10
±
3.22
14.47
±
2.11
11.06
±
5.49
Nodes on primary branch
20.83
±
1.40
22.37
±
3.19
15.77
±
2.09
18.55
±
3.50
Fertile nodes on primary branch
12.27
±
1.59
13.60
±
2.57
11.08
±
1.73
8.75
±
3.25
Nodes on secondary branch
16.52
±
1.52
21.52
±
2.70
13.27
±
2.01
14.04
±
3.32
Fertile nodes on secondary branch
8.47
±
1.43
12.33
±
2.71
7.75
±
1.07
5.63
±
2.15
Pod length (cm)
1.32
±
0.05
0.92
±
0.07
1.08
±
0.08
1.05
±
0.09
Pod breadth (cm)
0.60
±
0.00
0.50
±
0.00
0.54
±
0.05
0.54
±
0.06
Seeds pod-1
1.93
±
0.10
1.14
±
0.19
1.38
±
0.25
1.29
±
0.27
Pods plant-1
59.35
146.53
±
27.34
188.55
±
31.37
181.60
±
32.02
94.70
±
100 seed weight (g)
2.90
±
0.16
2.44
±
0.14
2.15
±
0.11
2.36
±
0.38
Biomass plant-1 (g)
12.97
±
2.90
11.53
±
2.54
10.86
±
3.36
5.37
±
4.18
Harvest index plant-1
42.83
±
2.99
35.63
±
5.07
28.20
±
5.32
33.69
±
8.57
Seed yield plant-1 (g)
5.56
±
1.32
4.06
±
0.88
2.94
±
0.83
1.90
±
1.69
235
Appendix XXXIII: Mean values and standard deviations for different plant traits in cross ILL 4605 x PL 406 Parameter
ILL 4605
Pant L 406
F1 generation
F2 generation
Days to flower
80.97
±
1.03
96.97
±
0.50
113.46
±
0.90
97.74
±
1.83
Days to mature
125.22
±
0.28
144.12
±
0.90
149.79
±
1.13
138.59
±
1.55
Plant height (cm)
26.05
±
0.31
31.50
±
1.05
34.14
±
0.56
24.14
±
1.18
Primary branches
2.57
±
0.15
2.60
±
0.10
2.63
±
0.16
2.12
±
0.17
Secondary branches
5.88
±
0.51
16.00
±
0.55
16.89
±
0.98
7.86
±
0.96
Nodes on primary branch
17.50
±
0.35
19.53
±
0.93
19.11
±
1.00
18.05
±
0.60
Fertile nodes on primary branch
6.53
±
0.28
11.70
±
1.02
11.26
±
0.68
7.65
±
0.70
Nodes on secondary branch
11.38
±
0.20
16.02
±
0.76
15.96
±
0.42
13.03
±
0.85
Fertile nodes on secondary branch
4.25
±
0.10
7.67
±
0.36
8.59
±
1.05
4.48
±
0.87
Pod length (cm)
1.31
±
0.01
1.00
±
0.02
1.05
±
0.02
1.05
±
0.02
Pod breadth (cm)
0.69
±
0.01
0.51
±
0.01
0.56
±
0.02
0.58
±
0.00
Seeds pod-1
1.33
±
0.06
1.94
±
0.03
1.29
±
0.07
1.18
±
0.05
Pods plant-1
35.40
±
0.98
145.15
±
14.04
148.65
±
7.21
43.12
±
11.07
100 seed weight (g)
3.94
±
0.04
1.86
±
0.01
1.98
±
0.01
2.20
±
0.03
Biomass plant-1 (g)
4.29
±
0.41
12.17
±
1.06
11.15
±
0.74
3.11
±
0.48
Harvest index plant-1
23.66
±
1.70
34.87
±
1.05
22.47
±
0.40
21.12
±
0.93
1.03
±
0.05
4.18
±
0.25
2.52
±
0.17
0.68
±
0.15
-1
Seed yield plant (g)
Appendix XXXIV: Mean values and standard deviations for different plant traits in cross PL 406 x ILL 4605 Parameter Days to flower
96.97
Pant L 406 ± 0.50
Days to mature
144.12
±
0.90
125.22
Plant height (cm)
31.50
±
1.05
Primary branches
2.60
±
0.10
Secondary branches
16.00
±
Nodes on primary branch
19.53
Fertile nodes on primary branch
11.70
Nodes on secondary branch
16.02
Fertile nodes on secondary branch
7.67
Pod length (cm)
1.00
Pod breadth (cm)
0.51
Seeds pod-1 Pods plant-1
80.97
ILL 4605 ±
F1 generation ± 6.12
97.55
F2 generation ± 18.44
1.03
113.17
±
0.28
149.48
±
2.67
139.40
±
14.11
26.05
±
0.31
35.00
±
2.61
24.86
±
6.23
2.57
±
0.15
2.70
±
0.56
2.14
±
0.64
0.55
5.88
±
0.51
16.48
±
4.16
7.86
±
4.08
±
0.93
17.50
±
0.35
18.37
±
3.61
16.00
±
3.71
±
1.02
6.53
±
0.28
11.52
±
1.79
6.80
±
2.52
±
0.76
11.38
±
0.20
15.97
±
3.59
13.26
±
4.41
±
0.36
4.25
±
0.10
8.78
±
2.31
4.96
±
2.36
±
0.02
1.31
±
0.01
1.09
±
0.07
1.04
±
0.09
±
0.01
0.69
±
0.01
0.59
±
0.06
0.56
±
0.06
1.94
±
0.03
1.33
±
0.06
1.33
±
0.18
1.30
±
0.34 34.07
145.15
±
14.04
35.40
±
0.98
154.07
±
28.81
46.85
±
100 seed weight (g)
1.86
±
0.01
3.94
±
0.04
1.98
±
0.22
2.31
±
0.40
Biomass plant-1 (g)
12.17
±
1.06
4.29
±
0.41
12.20
±
3.41
3.52
±
3.26
Harvest index plant-1
34.87
±
1.05
23.66
±
1.70
22.36
±
3.60
23.92
±
14.19
Seed yield plant-1 (g)
4.18
±
0.25
1.03
±
0.05
2.69
±
0.72
0.78
±
0.67
236
