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“EFFECT OF DROUGHT TOLERANT QTLs ON YIELD AND

YIELD CONTRIBUTING TRAITS UNDER MANAGED


WATER STRESS AS WELL AS NON STRESS CONDITION”

M.Sc. (Ag.) THESIS

By

MANOJ KUMAR NARANG

DEPARTMENT OF GENETICS AND PLANT BREEDING

COLLEGE OF AGRICULTURE
INDIRA GANDHI KRISHI VISHWAVIDYALAYA
RAIPUR (C.G.)

2011
“EFFECT OF DROUGHT TOLERANT QTLs ON YIELD AND
YIELD CONTRIBUTING TRAITS UNDER MANAGED
WATER STRESS AS WELL AS NON STRESS CONDITION”

Thesis

Submitted to the

Indira Gandhi Krishi Vishwavidyalaya, Raipur (C.G)

By

MANOJ KUMAR NARANG

IN PARTIAL FULFILMENT OF THE


REQUIREMENTS FOR THE
DEGREE OF

Master of Science

In

Agriculture
(GENETICS AND PLANT BREEDING)

Roll No. 12420 ID No. 110105044


JUNE 2011
CERTIFICATE-I

This is to certify that the thesis entitled “EFFECT OF DROUGHT

TOLERANT QTLs ON YIELD AND YIELD CONTRIBUTING TRAITS

UNDER MANAGED WATER STRESS AS WELL AS NON STRESS

CONDITION” submitted in partial fulfillment of the requirement for the degree

of “MASTER OF SCIENCE IN AGRICULTURE” of the Indira Gandhi

Krishi Vishwavidyalaya, Raipur, (C.G.) is a record of the bonafide research

work carried out by MANOJ KUMAR NARANG under my guidance and

supervision. The subject of the thesis has been approved by Student's Advisory

Committee and the Director of Instruction.

No part of the thesis has been submitted for any other degree or diploma or

has been published / published part has been fully acknowledged. All the

assistance and help received during the course of the investigation have been duly

acknowledged by her.

Date: Dr. S.B. Verulkar


(Chairman of the Advisory Committee)

THESIS APPROVED BY THE STUDENT’S ADVISORY COMMITTEE


Advisory Committee
Chairman : Dr. S.B. Verulkar _____________________

Member : Dr. A.K. Sarawgi _____________________

Member : Dr.(Smt.) Arti Guhey _____________________

Member : Dr. R.R. Saxena _____________________


CERTIFICATE – II

This is to certify that the thesis entitled “EFFECT OF DROUGHT

TOLERANT QTLs ON YIELD AND YIELD CONTRIBUTING TRAITS

UNDER MANAGED WATER STRESS AS WELL AS NON STRESS

CONDITION” submitted by Manoj Kumar Narang to the Indira Gandhi

Krishi Vishwavidyalaya, Raipur in partial fulfillment of the requirements for the

degree of M.Sc. (Ag.) in the Department of Genetics and Plant Breeding has

been approved by the External Examiner and Student‟s Advisory Committee after

oral examination.

EXTERNAL EXAMINER

Date:

Major Advisor ________________

Head of the Department ________________

Dean / Dean Faculty ________________

Director of Instructions ________________


ACKNOWLEDGEMENT

Research is not a single man show. It brings to light patience, vigor and dedication of
the person. It needs the close co-operation of the friends and colleagues and the guidance of
experts in the field to achieve something worthwhile.
Education plays fundamental role in personal and social development and teachers
play a fundamental role in imparting education. Teachers have crucial role in preparing young
people not only to face the further with confidence but also to build up it purpose and
responsibility. There is no substitute for teacher pupil relationship. A teacher is the one who
moulds our rough edges, to give us a smooth mind and you are the world of knowledge, love
and care. Once a year, we honor you all our lives we are grateful to say, you all my teachers.
"A journey is easier when you travel together; interdependence is certainly more
valuable than independence". I bow to Almighty “GOD” who keeps ever burning before my
vagrant steps the kindly light of hopes and always showered blessing on me without whose
endless benevolence and blessing this tedious task could not have been accomplished.
With a sense of high resolve and reverence, I would like to give my sincerest thanks to
my major advisor Professor (Dr.) S.B. Verulkar, Department of Genetics, and Plant Breeding
Indira Gandhi Krishi Vishwavidyalaya, Raipur (C.G.), for his precious instruction, extra
efforts, broad and profound knowledge, unique supervision, his patient instruction, sparing his
valuable time has given me a great inspiration and help at every step during my thesis and
pleasure to conduct this thesis under his supervision.
With a great reverence, I express my sincere thanks to respected members of my
advisory committee Dr. A.K. Sarawgi, Professor, Department of Genetics, and Plant Breeding
Dr. (Smt.) Arti Guhey, Professor, Department of Plant Physiology and Dr. R.R. Saxena,
Associate Professor, Department of Agricultural statistics and Social Science (language)
College of Agriculture, Raipur, for their critical suggestions, keen co-operation and kind help
rendered as and when needed.
I wish to record my sincere thanks to Dr. M.P. Pandey, Hon’ble Vice Chancellor, Dr.
S.K. Patil, Director of Research Services and Dr. U.K. Mishra, Director of Instructions,
IGKV, Raipur, for providing me the necessary facilities for research work.
Most humbly and respectfully I wish to express my profound sense of gratitude to Dr.
O.P. Kashyap, Dean, and Dr. R.L. Pandey, ExDean, College of Agriculture, IGKV, Raipur,
for his excellent guidance, valuable suggestion, memorable advices and encouragement which is
the vital source of inspiration in my life. I pay my sincere thanks to Dr. N.K. Motiramani,
Professor and Head, Department of Genetics and Plant Breeding for his excellence guidance
during course of investigation and providing necessary facilities.
I do express my heartily gratitude and indebtedness to Dr. (Smt) Ritu R. Saxena, Dr.
Rajeev Shrivastav, Dr. S.K. Nair and other respected teachers of the Department of genetics
and plant breeding for their wise counseling and encouragement during my research work.

Sincerest thanks are also extending to the all scientists of Department of the Genetics
and Plant Breeding, COA, Raipur, for their academic, technical and logistic help.
I would like to express my sincere gratitude to Dr. Madhav Pandey (Librarian, Nehru
Library, Raipur) and Shri U.K. Watti for giving me there kind help during my study.
I wish to express thanks to my seniors Rakesh Dhanwani, Sunil Verma, Naresh
Mahilange and Vikas Pali, Batch mates Amit, Kritraj, Manish, Madhuri, Prerna, Revati,
Roushan, Shivam, Shyam, Sujeet, Sunil, Tarun, Tikam, and Yogesh for their co-operation
during the course of work.
Unforgettable help and assistance throughout the course of this investigation by Mr.,
Makrand, Manish, Moti, Rajesh, Yuvraj,Mahesh and others also deserve my sincere thanks.

My words fail to express my deep sense of regards to my beloved parents, Shri P.R
.Narang (Babuji) Smt Meena Devi (Maaji). Mr. B. M. Narang (Dadaji),Usha Didi and Mr.
P.L.Johnson (Jijaji), Memin Didi, Thakurram (Bade Chachaji), Kajalkiran (Badi Chachiji),
Gaindram Narang (Manjhale Chachaji),, Harendra Narang (Chhote Chachaji), Mr. T. R.
Nirala (Fufaji), and other beloved family members whose love and silent blessings were the
source of my inspiration, encouragement and appreciation that opened the gate and pave the
way to my higher studies.

Indeed the words at my command are not adequate, either in form of spirit, to express
the depth of my humility and humbleness. Whenever I will open my thesis, each page of this
manuscript will tell story of contribution of my parents, advisor, seniors, colleagues, juniors
and well wishers. It will be untold strong and never be expresses in words.
Last but not the least; I would like to convey my cordial thanks to all those
unmentioned persons who helped me to fulfill my dream come true.

Department of Genetics and Plant Breeding

College of Agriculture, Manoj Kumar Narang


IGKV, Raipur (C.G.)
Dated:
CONTENTS

CHAPTER PARTICULAR PAGE NO.

I INTRODUCTION

II REVIEW OF LITERATURE

2.1 Impact of drought on yield and yield

contributing characters

2.2 Variability in rice for drought tolerance

2.3 Correlation of traits related to drought

tolerance

2.4 Mechanism of drought tolerance

2.5 Heritability of traits related to drought

tolerance

2.6 Genetic Variability

2.7 Correlation and Path Analysis

III MATERIALS AND METHODS

3.1 Location and climate

3.2 Experimental details

3.3 Methods

3.3.1 field studies

3.4 Observation

3.5 Statistical analysis

3.5.1 Analysis of variance

3.5.2 Parameters of variations


CHAPTER PARTICULAR PAGE NO.

3.6 Correlation coefficient analysis

3.7 Path analysis

IV RESULTS AND DISCUSSION

4.1 General observations

4.2 Variability

4.2.1 Analysis of variance

4.2.2 Drought Susceptibility Index

4.2.3 Mean and variability parameters

4.2.4 Genotypic and phenotypic coefficient of

variation

4.2.5 Heritability and genetic advance

4.3 Correlation studies

4.4 Path analysis

4.5 Effect of QTLs for Yield and yield

contributing traits

SUMMARY, CONCLUSION AND

SUGGESTIONS FOR FUTURE

RESEARCH WORK

ABSTRACT

REFERENCES

APPENDICES
LIST OF TABLES

TABLE PARTICULARS PAGE


NO. NO.
3.1 Details of the breeding materials
4.1 Analysis of variance for different characters in rice under
terminal stage drought and irrigated condition during
Kharif 2010
4.2 Mean of grain yield with Drought Susceptibility Index in
Terminal Stage Drought and Irrigated condition
4.3 Genetics parameters of variation for yield and its
components under TSD and Irrigated condition (Kharif
2010)
4.4 Correlation coefficient of grain yield and its component in
with QTL DTY 1.1 line
4.5 Correlation coefficient of grain yield and its component in
with QTL DTY 1.1 & 3.2 line
4.6 Correlation coefficient of grain yield and its component in
with QTL DTY 3.2 lines
4.7 Correlation coefficient of grain yield and its component in
without QTL lines
4.8 Correlation coefficient of grain yield and its component in
check lines
4.9 Path coefficient analysis showing direct and indirect
effects of different yield contributing character on yield in
with QTL DTY 1.1 lines
4.10 Path coefficient analysis showing direct and indirect
effects of different yield contributing character on yield in
with QTL DTY 1.1 & 3.2 lines
4.11 Path coefficient analysis showing direct and indirect
effects of different yield contributing character on yield in
with QTL DTY 3.2 lines
TABLE PARTICULARS PAGE
NO.
NO.
4.12 Path coefficient analysis showing direct and indirect
effects of different yield contributing character on yield in
without QTL lines
4.13 Mean performance and percent change in days to 50%
flowering in lines with QTL and without QTL lines
under Terminal stage drought and irrigated condition.
4.14 Mean performance and percent change in Plant height in
lines with QTL and without QTL lines under Terminal
stage drought and irrigated condition
4.15 Mean performance and percent change in Panicle length
in lines with QTL and without QTL lines under Terminal
stage drought and irrigated condition.
4.16 Mean performance and percent change in Flag leaf length
in lines with QTL and without QTL lines under Terminal
stage drought and irrigated condition.
4.17 Mean performance and percent change in Flag leaf width
in lines with QTL and without QTL lines under Terminal
stage drought and irrigated condition.
4.18 Mean performance and percent change in flag leaf area in
lines with QTL and without QTL lines under Terminal
stage drought and irrigated condition.
4.19 Mean performance and percent change in Grain length in
lines with QTL and without QTL lines under Terminal
stage drought and irrigated condition.
4.20 Mean performance and percent change in Grain width in
lines with QTL and without QTL lines under Terminal
stage drought and irrigated condition.
4.21 Mean performance and percent change in grain L: B ratio
in lines with QTL and without QTL lines under Terminal
stage drought and irrigated condition.
TABLE PARTICULARS PAGE
NO. NO.

4.22 Mean performance and percent change in Seed Index in


lines with QTL and without QTL lines under Terminal
stage drought and irrigated condition.
4.23 Mean performance and percent change in Harvest Index
in lines with QTL and without QTL lines under Terminal
stage drought and irrigated condition.
4.24 Mean performance and percent change in Biological yield
in lines with QTL and without QTL lines under Terminal
stage drought and irrigated condition.
4.25 Mean performance and percent change in Grain yield in
lines with QTL and without QTL lines under terminal
stage drought and irrigated condition.
LIST OF FIGURES

Figure Particulars Between pages


no.
3.1 Distribution of rainfall during wet season 2010

LIST OF APPENDICES

Appendix Particulars Page No.

I Weekly meteorological data during crop growth

period of (Kharif 2010)

II List of genotypes

CHAPTER-I

INTRODUCTION

Rice is the most consumed cereal grain in the world, constituting the

dietary staple food for more than half of the planet‟s human population. Apart

from food rice is intimately involved in the culture as well as economy of many

societies. Rice is an integral part of creation myth and remains today as leading

crop and most preferred food (Huke and Huke, 1997). Possibly the oldest

domesticated grain (~10000 years), rice is the staple food for 3 billion people (60%

of the world‟s population) and growing rice is the largest single use of land for
producing food, covering 9% of the earth's arable land. Rice a member of the

family Poaceae originated from South-East Asia and in the Asia, where more than

90% of world‟s rice is produced and consumed (Li and Xu, 2007) thus rice is

immensely important to food security of Asia. Out of 24 species only two species

(O. glaberima and O. sativa) are cultivated.

In world, rice has occupied an area of 154.32 million hectares, with a total

production of 661.8 million tons in 2008 (Anonymous, 2009). In Asian countries,

rice is the main staple crop covering about ninety per cent of rice grown in the

world, with two countries, China and India, growing more than half of the total

crop. Rice provides about two-third of the calorie intake for more than two billion

people in Asia, and a third of the calorie intake of nearly one billion people in

Africa and Latin America (Shastry et al., 2000)

India, the second largest producer of rice after China has an area of over

43.77 million hectares with the production of 96.43 million tons in 2008

(Anonymous, 2009). Rice being the main source of livelihood for more than 120 –

150 million rural household is the backbone of the Indian Agriculture. It occupies

about 23.3 per cent of the food grain production and 55 percent of cereal

production. The rice plays a vital role in the national food security. Even then rice

self-sufficiency in India is precarious. The country's population of more than a

billion is growing at 1.8% per year, outpacing the 1.4% annual growth rate of rice

production.

In India, rice is grown under four ecosystems: irrigated, rainfed lowland,

rainfed upland and flood prone. More than half of the rice area (55%) is rainfed

and distribution wise 80% of the rainfed rice areas are in eastern India, making its
cultivation vulnerable to vagaries of monsoon. Even through the region receives

good rainfall yield losses caused by drought every year reach 2.9 million tons

annually (Widawsky and O‟Toole, 1990).

Chhattisgarh popularly known as “Rice Bowl of India” occupies an area

around 3610.47 thousand hectares with the production of 5.48 MT and

productivity of 1517 kg ha-1 (Anonymous, 2010). The prime causes of low

productivity of rice in Chhattisgarh are inappropriate adoption of agronomical

practices, limited irrigation (28.0 %), lack of improved varieties suitable to

different ecosystems and lack of extension services. Chhattisgarh farmers are

mainly depending on climate for rice cultivation.

In general, rice is grown in the diverse environment conditions of a wide

range of latitude and altitudes. The rice crop need hot and humid climate. It is best

suited to the regions which have high humidity, prolonged sunshine and an assured

supply of water.

The major climatic factor affecting growth and yield include solar

radiation, temperature and rainfall (the later is particular important in rainfed rice

culture). Temperature, solar radiation and rainfall are important weather factor that

influence rice yield directly by affecting the physiological process involved in

grain production and indirectly, through their effect on disease and insect pressure.

These factors are often difficult to differentiate in the field. In temperate regions,

the rice calendar is circumscribed by the temperature regime. In the tropics, it is

determined primarily by onset and withdrawal of the monsoon (Seshu and Caddy,

1989). The average requirement for blooming is in the range of 26.50C to 29.50C
while at the time of ripening the temperature should be in between 20-250C (De

Datta, 1981).

Drought stress is a major constraint to rice production and yield stability in

rainfed region (Evenson et al., 1996). It acts as a serious limiting factor in

agriculture production by preventing a crop from reaching the genetically

determined theoretical maximum yield. The effect of drought on crop production

and overall economy is well known. The annual reduction in rice yields due to

drought averages 18 million tons globally (O‟ Toole, 2004). In Asia alone, it is

estimated that a total of 20 million ha of rice field (16 million ha in upland and 13

million ha in lowland) are drought prone. In the eastern states of India viz.

Jharkhand, Orissa and C.G. alone, total rice production losses in severe droughts

(about 1year in 5) averages 40%, valued at $ 650 million (Pandey et al., 2005). In

these areas, drought risk reduces productivity even in non drought years, because

farmers avoid purchasing inputs when they fear crop loss, becoming mired in cycle

of low productivity, poverty and food insecurity (Pandey et al., 2005). Rice is

particularly sensitive to drought stress during reproductive growth, when even

moderate stress can result in drastic reduction in grain yield. The development of

cultivars with improved drought tolerance is thus an important element in reducing

risk; increasing productivity and alleviating poverty in communities depend on

rainfed production.

Compared with other cereals, rice requires more water to produce good

yields. However, declining water supplies are compelling rice researchers to

explore new ways for producing high yields with less water (Farooq et al., 2010).

An approach to minimizing agricultural losses incurred by drought stress is to


develop, via, genetic means, plant cultivars that can escape or withstand periods of

drought. Such development should have a lasting economic impact on agriculture

worldwide. However, despite many decades of research, drought tolerance

continues to a major challenge to plant breeders, partly because of the perceived

complexity of the traits. Accumulating evidence suggests that plant response to

drought tolerance is controlled by more than one gene (Zhang et al., 2001) and is

highly influenced by environmental variation. The quantification of drought

tolerance poses serious difficulties. Direct selection in the field is not always

possible because uncontrollable environmental factors, such as variation in

rainfall, interactions with extreme temperatures and variations in salinity and

nutrient availability adversely affect the precision and repeatability of such traits

(Nguyen et al., 1997).

Since drought is stage specific phenomenon and is developmentally

regulated, selection and breeding is difficult. Tolerance at one stage of plant

development may be poorly correlated with tolerance at other developmental

stages. Specific stages throughout the ontogeny of the plant, including seed

germination and emergence, seedling survival and growth, vegetative and

reproductive growth should be evaluated separately for the assessment of drought

tolerance and identification of genetic components. Each developmental stage,

which should be considered as a separate trait, may require a different screening

procedure and simultaneous or sequential screening may be impractical or

impossible. Furthermore, selection for drought tolerance using phenotypic

measurements requires specialized personnel and extensive investment in field

nurseries or green house facilities. These complexities have led to limited success
in developing drought-tolerant plants or improving crop yields in dry

environments.

Despite the importance of drought as constraints, little efforts have been

devoted to developing drought tolerant rice cultivars. Most improved cultivars

grown in drought prone area are originally bred for irrigated conditions and were

not selected for drought tolerance. These cultivars have high yield potential but

often highly prone to yield reduction under drought. On the other hand some

traditional rice cultivars grown in rainfed areas are highly drought tolerant but

have low yield potential. In order to have high yield in non drought years, farmers

require cultivars that combine high yield potential with improved tolerance to

drought (Kumar et al., 2008).

Progress is being made in developing drought tolerant rice germplasm

through conventional breeding (physiological dissection) and the use of molecular

tools. Identification of quantitative trait loci (QTL) conferring improved drought

resistance may facilitate breeding progress (Bernier et al., 2009a). The advent of

saturated molecular maps promised rapid progress towards the improvement of

crops for genetically complex traits like drought resistance via. analysis of

quantitative trait loci (QTL) (Price et al., 2002a).

Many important agronomic traits in crop plants, including stress tolerance,

are complex traits controlled by quantitative trait loci (QTLs). Isolation of these

QTLs holds great promise to improve world agriculture but is a challenging task

(Zhong Hai et al. 2005).

In QTL analysis, rice has been receiving exceptional attention, because it

has been used as a model plant in molecular biology. It contains approximately 3.5
times as much DNA as Arabidopsis, but only approximately 20% as much as

maize and approximately 3% as much DNA as wheat. In this chapter, QTL will be

viewed globally from different perspectives, using rice as a model (Xu, 2001).

QTLs are marvelous genetic entities demonstrating great utility in genetic

understanding of complex traits and promise greater impact in crop improvement

endeavors they have given as a clearer view of the genetics of agronomically

important and several other traits too. The distinction and clarity between

monogenic and polygenic traits is blurred with the increase in power and precision

of methods to detect and map QTLs (Shashidhar, 2005).

Keeping the above points in view, the present piece of research was

undertaken with the following objectives:

1 To understand the effect of DT QTLs on yield under stress condition.

2. To understand the effect of DT QTLs on yield under non-stress condition.


CHAPTER II

REVIEW OF LITERATURE

Drought is the most important abiotic stress in rainfed. Approximately 45%

of world rice is grown under rainfed condition (Verulkar et al 2010). The

estimated average loss in rice production during drought years for eastern India is

5.4 million tones. In Asia alone, about 34 million ha of rainfed lowland and 8

million ha of upland rice (Huke and Huke, 1997) are subject to frequent drought

stress. In this area water shortage is the major constraint in productivity, and for

this reason drought resistance has been targeted by breeders (Nguyen et al., 1997).

One of the limitations in developing drought tolerant genotypes in rice is

narrow genetic base and most of the known drought tolerant donors have low

breeding value and progenies are low yielding particularly under non-stress

condition.

Achieving drought resistance in rice will be necessary for meeting the

growing water shortage of the world. And it requires a deeper understanding of the

mechanism that could facilitate drought resistance (Serraj et al., 2011). Progress is

being made in developing drought tolerant rice germplasm through conventional

breeding (physiological dissection) and the use of molecular tools. Identification of

quantitative trait loci (QTL) conferring improved drought resistance may facilitate

breeding progress (Bernier et al., 2009). Recently, QTLs with large effect on grain

yield under drought stress (DT QTLs) has been indentified along with

development of lines with and without QTLs. This study aims to understand the

effect of these DT QTLs on yield and yield contributing traits under managed
water stress as well as non stress condition. The available literature has been

classified into following headings:

2.1. Impact of drought on yield and yield contributing characters

2.2 Variability in rice for drought tolerance

2.3. Correlation of traits related to drought tolerance

2.4. Mechanism of drought tolerance

2.5. Heritability of traits related to drought tolerance

2.6. Genetic Variability

2.7. Correlation and Path Analysis

2.1. Impact of drought on yield and yield contributing characters

Puckridge and O‟Toole (1981) shown that biomass production of rice is a

function of water use. The shortage of water in the soil suppresses the leaf

expansion, tillering and photosynthetic rate along with leaf area due to

senescence. All these factors are responsible for reduction in dry matter

accumulation.

Row et al. (1983) studied the yield component of rice varieties viz. Rasi

and IR-20 under moisture stress condition. The reproductive and ripening phase

were vulnerable and crucial for moisture stress, which resulted in permanent

damage to growth and yield.

Krishnaya and Murty (1991) exposed five upland rice varieties to soil

moisture stress of 25% field capacity at seedling stage. With increase in soil

moisture stress, there was a decrease in stress, relative water content and water

potential of leaf. The cultivar which maintained high relative water content and
positive turgor, in spite of reduced leaf water potential during stress also had

optimum photosynthesis and solute accumulation.

Woperesis et al. (1996) reported that water stress during flower induction

and inflorescence development leads to delay in flowering (anthesis) or even to

complete inhibition as apical morphogenesis is sensitive to water deficit.

Pantuwan et al. (2002) reported that the drought stress developed prior to

flowering generally delayed the flowering of genotypes and such a delay was

associated with drought susceptibility in rice.

Deming et al. (2004) found that drought stress during reproductive stages

showed severe drought stress in panicle development stage caused great yield

loss by reducing spikelet per panicle in upland rice.

Deshmukh et al. (2007) observed that in rice, water stress during flowering

can reduce the harvest index by as much as 60%, largely as a result of a reduction

in grain set. Panicles in stress plants fail to full exert (emerge) from the flag leaf

sheath, flowering is delayed and the percentage of spikelet that open at anthesis is

reduced.

Large scale screening for drought tolerance in field is complicated because

of difference in phenology and rains during drought stress periods (Blum, 2002).

Drought resistance traits are divided into primary traits, secondary traits,

integrative traits, phenology, and plant-type traits. Primary traits are further

divided into constitutive traits (i.e. under no drought stress; e.g. rooting depth, root

thickness, branching angle and root distribution pattern (Lafitte et al., 2001) and

induced traits (e.g. hardpan penetration and osmotic adjustment). Secondary traits

include maintenance of plant water status, canopy temperature, leaf rolling score
and leaf death score. Secondary traits are also useful for screening (Lafitte et al.,

2004).

Transfer of primary or secondary traits such as those associated with root

growth to desirable background to enhance grain yield is complicated by a lack of

clear understanding of the genetic of component traits and their interaction and a

lack of tightly linked marker Uga et al (2011).

Although yield under stress generally have higher phenotypic correlation

with some of yield component (e.g. grain number and percent fertile spikelets;

integrative traits) then they do with primary traits, primary traits are likely to be

controlled by fewer underlying gene or QTLs. Therefore molecular

characterization of primary traits (e.g. QTL analysis) will presumably be a more

promising avenue then the study of yield components.

Most of the improved rice cultivars now grown in the rainfed drought-

prone areas were bred for irrigated conditions and are highly susceptible to

drought. Recently, QTLs with large effects on grain yield under drought stress,

namely DTY1.1, DTY2.1, DTY2.2, DTY3.1, DTY4.1, DTY9.1 and DTY12.1 have been

identified by IRRI scientists, explaining 31-77% of the phenotypic variance.

Molecular approaches to drought tolerance have been widely applied to

rice, beginning with QTL analysis. Several experiments have been reported for

yield and yield component under different type of drought stress in rice. QTL have

been detected for several root related traits and osmotic adjustment (OA) in rice

(Li et al., 2005). The Way Rarem-derived allele of qtl12.1 was confirmed to

improve grain yield under drought mainly through a slight improvement (7%) in

plant water uptake under water-limited conditions (Bernier et al., 2009).


2.2 Variability in rice for drought tolerance

Success in crop improvement depends on the magnitude of genetic

variability and the extent to which the desirable characters are heritable. The work

on estimates of genetic variability has been carried out earlier in rice, however

scarce information is available on these aspects in upland rice.

Singh et al. (1984) reported grain yield/ m2 had maximum GCV followed

by plant height. However, Rahangdale and Khorgade (1988) also reported higher

GCV for grain yield.

Husain et al. (1987) reported high genotypic coefficient of variations for

grain length: breadth ratio and grain breadth.

Datta and Rao (1988) studied Genetic variability of plant characters in

thirty rice strains and founded high genetic coefficient of variation was observed

for grain yield/ m2

Genetic variation has been observed in rice for plasticity and also in

response to soil drying after flooding Suralta et al.( 2010 )

Veeresh et. al (2011).predicted that variation in rice root response to

drought from a physiological perspective in terms of morphology and function

with respect to the different growth environments

Genetic differences have also been observed in the ability of seminal roots

to continue elongation and form parenchyma under flooded conditions after

drought.( Suralta and Yamauchi 2008; Suralta et al.,2008a,b,2010)

Different genotype have exhibited different responses in plasticity

depending on type of drought stress.( Kano et al., 2011)


Chauhan et al. (1993) studied 21 advanced breeding lines/ cultivars and

reported that high expected genetic advance coupled with high heritability was

observed for days to 50% flowering and panicle length and low to moderate

heritability for grain yield.

Patil et al. (1993) studied 37 promising cultures of upland rice. They

reported high genotypic coefficient of variation for grain yield/ m2 and plant

height.

Chaudhary and Das (1998) worked out estimation of variability in 11 deep

water rice varieties and its attributing characters. High genotypic coefficient of

variation was observed for grain yield.

Singh et al. (1999) studied F5 generation of rice cross genotypic coefficient

of variation and phenotypic coefficient of variation higher for all the character

except for days to 50% flowering.

The high GCV estimates for number of productive tillers per plant, number

of spikelet per panicle and grain yield per plant was reported by Shivani and

Reddy (2000), Gomez and Rangasamy (2002) and Gomez and Kalamani (2003).

These traits showing high genotypic variability offer greater scope for genetic

improvement through selection.

Umadevi et al. (2009) reported that PCV was higher than GCV for all the

characters studied. PCV ranged from days to 50% flowering to leaf length. GCV

and PCV was low (< 10%) for days to 50% flowering and grain length. Similar

findings of low GCV (7.14%) were reported earlier for days to 50 per cent

flowering (Shivani and Reddy, 2000). GCV for plant height, panicle length, leaf
breadth, 100 grain weight, kernel breadth, kernel length/ breadth ratio and grain

yield per plant recorded moderate (10-20%) level of GCV indicating considerable

amount of variability expressed for these characters and high GCV (> 20%) for

total number of productive tillers per plant (20.01%), number of secondary

branches, leaf length and straw yield.

2.3 Correlation of traits related to drought tolerance

Shahid et al. (1994) reported positive correlation between root length, root

dry weight, shoot dry weight, stomata frequency and drought tolerance, whereas

negative correlation existed between shoot length and stomata size.

Babu et al. (2003) reported positive correlation of biomass under stress

with yield, per cent spikelet fertility, number of grains per panicles, harvest index

and relative yield. On the other hand, leaf rolling scores had negative correlation

with yield and harvest index.

Kumar et al. (2004) observed that in rice there is a negative association

between delay in flowering and grain yield, relative water content and post

flowering dry matter production under rainfed condition. A direct relationship was

observed between delay in flowering and sterility. The flowering delay under

drought stress condition is governed by leaf water status.

Lafitte et al. (2004) found negative correlation between plant height and

yield. Tall lines yield poorly in rainfed experiment but no significant association

was observed between yield and height under control condition.

Lanceras et al. (2004) using double haploid mapping population measured

correlation between traits and grain yield at different water levels reported

correlation between grain yield and biological yield ranges 0.70 to 0.54, indicating
that genetic improvement in the grain yield would likely be accompanied by

improvement of biological yield. Further they found that with increase drought

stress, correlation between grain yield and harvest index increase dramatically,

indicating that harvest index is also a primary determinant of grain yield. Positive

correlation was also found between total spikelet number and grain yield under

stress condition. Negative correlation was found between grain yield and days to

flowering after initiation of irrigation gradient.

Singh et al. (2004) reported significant and positive correlation between

grain yield per plant and yield contributing traits, effective flag leaf breadth and

total grains per panicle. A negative correlation was found between grain yield and

plant height.

Verulkar et al. (2004) reported positive correlation of spikelet fertility,

harvest index and biological yield with yield under stress.

Zou et al. (2005) found significant positive correlation between canopy

temperature and spikelet fertility, leaf water potential, grain weight per plant and

leaf rolling score in stress. However, there was a significant negative correlation

only between the canopy temperature and spikelet fertility, grain weight per

panicle under normal conditions.

Gomez et al. (2005) observed that biological yield and plant height was

positively correlated with grain number/ panicle. Leaf area per plant showed the

highest positive direct effect on root weight, followed by biological yield and root

breadth. Selection based on biological yield may help identify drought-resistant

types.
Gomez et al. (2006) reported positive and significant correlation of panicle

length and number of productive tillers with grain yield under stress. They also

observed positive correlation of root traits with plant height under stress. Similar

relation was also reported earlier in rice by Venuprasad et al. (2002).

Umadevi et al. (2009) reported that grain yield per plant was positively and

significantly associated with days to 50% flowering, number of productive tillers

per plant, number of secondary branches per panicle and straw yield.

Fu et al. (2010) reported correlations of grain yield per plant with spikelet

per panicle, 1000 grain weight, number of panicles per plant and percentage seed

set was significant positive. The number of grains per panicle also had strong

positive correlations with spikelet per panicle and percentage seed set, but the

number of panicles per plant had significant negative correlations with spikelet per

panicle, grains per panicle and 1000 grain weight. In addition, there was a

significant negative correlation between panicles per plant and percentage seed set.

Gomez et al. (2010) reported significant positive correlation between plant

height and biomass under stress. Significant negative correlations were noticed

between water stress indicators such as leaf rolling and leaf drying with biomass

under stress.

Negative correlations of water stress indicators with biomass and positive

correlations of plant production traits with biomass have been reported in rice. It

was also reported by Babu et al. (2003), Gomez et al. (2006) and Srinivasan et al.

(2008).

2.4 Mechanism of drought tolerance


Rice, like other crops can potentially resist drought stress using three

different strategies: drought escape, drought avoidance and drought tolerance

(Bernier et al., 2008). A proper timing of life cycle, resulting in the completion of

most sensitive developmental stages while water is abundant, is considered to be a

drought escape strategy (Price et al., 2002b). This mechanism involves rapid

phonological development (early flowering and early maturity), developmental

plasticity (variation in growth period depending on extend of water-deficit) and

remobilization of pre-anthesis assimilates to grain.

Drought avoidance is the ability of the plant to maintain relatively high

tissue water potential despite a shortage of moisture. Mechanisms for improving

water uptake, storing in plant cell and reducing water losses confer drought

avoidance. Drought avoidance is performed by maintenance of turgor through

increased rooting depth, efficient root system and increased hydraulic conductance

and by reduction of water loss through reduced epidermal (stomatal and lenticular)

conductance, reduced absorption of radiation by leaf rolling or folding and reduced

evaporation surface (leaf area) (Passioura, 1976). Number of worker have

emphasized the importance of traits responsible for avoidance mechanism of

drought tolerance.

Drought affects rice at morphological, physiological and molecular levels

such as delayed flowering, reduced dry matter accumulation and partitioning and

decreased photosynthetic capacity as a result of stomatal closure, metabolic

limitations and oxidative damage to chloroplasts. Small-statured rice plants with

reduced leaf area and short growth duration are better able to tolerate drought

stress, although the mechanisms are not yet fully understood. Increased water
uptake by developing larger and deeper root systems, and the accumulation of

osmolytes and osmoprotectants are other important mechanisms for drought

resistance (Farooq et al., 2009).

Mechanism such as osmotic adjustment where by a plant maintains cell

turgor pressure under reduced soil water potential, increase in elasticity in cell and

decrease in cell size and desiccation tolerances by protoplast are categorized as

drought tolerance mechanisms (Sullivan and Ross, 1978).

Veeresh et. al (2011)temporal and spatial heterogeneity is a possible path

toward the use of root and dehydration avoidance traits for improving drought

resistance in rice

O‟Toole and Datta (1986) suggested that increase root depth and root

density in rice increase the capacity to extract available soil water and may be

responsible for increase drought avoidance in some rice genotypes.

Hacke et al. (2001) and Stiller et al. (2003) revealed that the wider xylem

vessels may decrease chances of cavitations, which would reduce xylem water

conductivity.

However, Passioura (2006) reported that the contribution of root depth to

drought avoidance may be site specific, as the subsoil is often inhospitable to root

growth due to unfavorable pH or nutrient content.

2.5 Heritability of traits related to drought tolerance


Babu et al. (2003) reported high heritability (broad sense) of leaf rolling,

grain yield, plant height, spikelet fertility, harvest index and days to heading (0.65,

0.59, 0.49, 0.43, 0.60 and 0.70 respectively), while low to moderate for relative

water content, plant height, grain yield, biomass, spikelet fertility, grain per

panicle and 1000 grain weight under stress.

Lanceras et al. (2004) computed broad sense heritability (h2) across five

water regimes using 154 double haploid mapping population derived from a cross

between two rice cultivars, CT9993-510 to 1-M and IR62266-42 to 6-2. They

reported that panicle number and plant height had highest h2, 0.61 and 0.73,

respectively, harvest index and spikelet fertility had moderate h2, 0.46 and 0.45

respectively, while grain yield and biological yield had the lowest h2, 0.32 and

0.31 respectively.

Singh et al. (2004) observed high heritability of traits like grain yield and

yield contributing character, suggesting that the importance of these traits as

criteria for improving overall yield under stress. Moderate heritability of panicle

length and low heritability of leaf breadth and test weight.

Kumar et al. (2008) reported high heritability ( broad sense) for grain yield,

harvest index, 50% flowering, leaf rolling date and leaf rolling was 0.48, 0.45,

0.47, 0.20 and 0.42, respectively for IR55419/ IR64 and 0.45, 0.38, 0.73, 0.26 and

0.30, respectively for IR55419/ Way Rarem.

Venuprasad et al. (2009) reported heritability in broad sense under lowland

stress was 0.73 in 2006 and 0.55 in 2007 for grain yield.
Bernier et al. (2009) reported heritability 0.75, 0.49, 0.54, 0.94 and 0.44 for

grain yield, biomass yield, harvest index, days to 50% flowering and plant height,

respectively.

Umadevi et al. (2009) estimated heritability in broad sense and genetic

advance as percentage of mean of days to 50% flowering 0.93 and 14.25, plant

height (cm) 0.99 and 28.54, panicle length (cm) 0.87 and 20.26, number of

secondary branches/ panicle 0.91 and 39.03, leaf length (cm) 0.99 and 61.87, leaf

width (cm) 0.92 and 27.34, grain length (mm) 0.98 and 17.68, grain breadth (mm)

0.94 and 22.82, grain L/ B ratio 0.95 and 33.19, straw yield (g) 0.99 and 58.37,

100 grain weight (g) 0.93 and 28.95, and grain yield per plant (g) 0.78 and 29.36

respectively.

Verulkar et al. (2010) reported broad sense heritability of 50% flowering

(0.92 for control, 0.78 for moderate and 0.83 for severe drought), plant height

(0.96 for control, 0.81 for moderate and 0.83 for severe drought), grain yield (0.55

for control, 0.52 for moderate and 0.73 for severe drought) and harvest index (0.58

for control, 0.56 for moderate and 0.78 for severe drought).

2.6 Genetic Variability

The basic requirement of plant breeding is the presence of genetic

variability. Importance of genetic variability in any breeding material is a

prerequisite as it provides not only a basis for selection but also some valuable

information regarding selection of diverse parents for use in hybridization

programme. Variability in rice, before it became a subject of study by plant

breeder was the concern of taxonomists such as Linnaeus (who gave the generic
name „Oryza‟), De Candolle (1886), Camus (1920), Roschevicz (1932), Chatterjee

(1947) and Porter (1956).

2.6.1 PCV and GCV

Verma et al. (2000) reported that the higher magnitude of genotypic

coefficient of variation and phenotypic coefficient of variation were observed for

number of sterile and total spikelets per panicle, grain yield, biological yield,

number of productive and total tillers per plant.

Iftekharuddaula et al. (2001) evaluated nineteen rice hybrids in order to

determine variability and it was observed that all the tested character had

significant variation. The highest genotypic coefficient of variation for panicles

per plant followed by spikelets per panicles and grain yield.

Das et al. (2001) investigated 29 boro rice genotypes to study genotypic

and phenotypic coefficient of variation, broad sense heritability and genetic

advance for plant height, number of panicles plant-1, panicle length, number of

filled grains panicle-1, 100-grain weight, grain yield and harvest index. For all the

characters, the estimates of GCV were smaller than that of PCV, suggesting the

influence of environment on them. High heritability coupled with high genetic

advance for plant height, panicles plant-1, harvest index and grain yield plant-
1
indicated that these characters were predominantly governed by additive gene

action. High heritability and moderate to low genetic advance for panicle length,

filled grains panicle-1 and 100-grain weight suggested the predominant role of non-

additive gene action in their inheritance.

Singh et al. (2002) evaluated the phenotypic and genotypic variability in

fifteen quantitative characters of 52 genotypes of lowland rice. The genotypic and


phenotypic variances were high for grain yield per plant, panicle weight, number

of grains per panicle and number of branches per panicle, grains per panicle and

100 seed weight and low for panicle length. Heritability in the broad sense ranged

from 3.61 for number of effective tillers per plant to 99.57 for grain length. High

heritability with high genetic advance was recorded for number of grains per

panicle followed by panicle weight and grain yield per plant.

Bhandarkar et al. (2002) worked out estimates of variability, genetic

advance as percentage of mean in 22 semi deep-water rice genotypes for yield and

its attributing characters. High heritability with high genetic advance as percent of

mean were observed for panicle weight, significant negative correlations were

observed for days to 50% flowering, maturity, plant height with grain yield per

plant.

Kavitha and Reddy (2002) studied variability and heritability in twenty one

genotypes of rice. Considerable amount of variability was observed for all the

quantitative characters studied.

Gomez and Kalamani (2003) conducted a study for physiological and

economic traits (Plant height, number of productive tillers per plant, panical

length, 1000 grain weight, biological yield per plant) in 11 rice landraces and

reported significant variation among land racers for all characters, especially

biological yield per plant.

Khedikar et al. (2003) studied genetic variability for seven characters viz.

days to 50% flowering, plant height, effective tillers per plant, panicle length, test

weight, head rice recovery and grain yield per plant in 20 scented rice genotypes

grown under 3 different environments. The PCV was higher than GCV among all
characters, indicating that the variation among them is not only due to genetic

content but also due to the influence of environment.

Mishra et al. (2003) evaluated 88 rice genotypes and reported presence of

considerable amount of variability for the plant height, flag leaf width, ear bearing

tillers per plant, 100-seed weight, hulling and milling percentage, panicle length,

biological yield, harvest index, kernel length after cooking, gelatinization

temperature and grain yield. The high genotypic variation was observed for

biological yield and grain yield per plant.

Sarma and Bhuyan (2004) studied the genetic variation in fifty eight ahu

rice genotypes, consisting mostly of indigenous and improved cultivars in Assam,

India during the summer season of 2000. They used grain yield per plant and 8

yield components for estimating the genetic parameters and in the D2 analysis.

They reported highest PCV and GCV being exhibited by the number of grains per

panicle, followed grain yield per plant and number of effective panicles per plant.

Medhi et al. (2004) studied forty indigenous scented rice varieties of

Assam including an introduction from Thailand and indicated that there was

considerable variation among the varieties for all the traits. The estimates of

genotypic and phenotypic coefficient of variation were found to be high for grain

per panicle, 100-grain weight, grain yield per plant and effective tillers.

Souroush et al. (2004) evaluated 36 genotypes and determined the

relationship between grain yield and yield components. Analysis of variance

showed a highly significant difference among the genotypes for all the traits.

Tyagi et al. (2004) evaluated seventy rice genotypes for genetic variability

and association among some seedlings and mature plant characters. Significant
genotypic variations were observed for most of the character studied. The yield

components exhibited high heritability coupled with high genetic advance.

Singh et al. (2005) studied seven characters in twenty rice genotypes

including a local check during Kharif 2004. Through analysis of variance, they

reported significant differences among the genotypes for all the characters. High

estimates of (GCV) and (PCV) were reported for grain yield, biological yield,

grain density and harvest index (%).

Singh and Singh (2005) working with twenty promising mutants of Pusa

Basmati-1 rice reported high PCV and GCV for head rice recovery, aroma and

alkali digestion value; which suggest that there is a possibility of improving these

characters through direct selection.

Hasib (2005) reported high phenotypic and genotypic coefficient of

variation, heritability and genetic advance for spikelets number per panicle, test

weight and grain yield per panicle indicating important role of additive gene action

for the expression of these traits.

Bisne et al. (2006) observed high GCV for filled spikelets per panicle,

chaffy spikelets per panicle, spikelet fertility percentage, 100 grain weight, harvest

index and grain yield per plant.

Veni et al. (2006) studied variability and heritability for 7 important

physio-chemical traits viz., kernel length, kernel breadth, length breadth ratio,

kernel length after cooking, elongation ratio and alkali spreading value and

reported low to moderate estimates of variability, moderate to high heritability and

low expected genetic advance for all the characters indicated the preponderance of

both additive and non-additive gene effects in conditioning these traits.


Devi et al. (2006) conducted a study on fifty four standard rice genotypes

that are well adapted to different climatic conditions. They reported higher PCV

than GCV for all the characters plant height, panicle length, tillers per hill, 50%

flowering, maturity duration and grain yield per plant.

Monalisa et al. (2006) studied twenty lowland rice cultivars of West

Bengal. They reported highest GCV for high density grains per panicle and panicle

weight.

Jaiswal et al. (2007) evaluated twenty five aromatic rice genotypes for

yield and quality characters to assess the genetic variability. Highest GCV was

recorded for grain yield per plant and number of panicle bearing tillers. High

heritability coupled with high genetic advance was observed for yield per plant,

number of panicle bearing tillers and number of grains per panicle.

Sharma et al. (2008) reported significant differences among rice cultures

for milling and physico-chemical traits.

Bisne et al. (2009) found that low, moderate, and high genotypic and

phenotypic coefficient of variation were expressed by harvest index, total number

of filled spikelet‟s per panicle, 100-grain weight and spikelet fertility percentage.

Patil et al. (2009) conclude that genotype exhibited significant variability

for days to 50% flowering, flag leaf length, plant height, panicle length, number of

filled grains per panicle, 100 seed weight and grain yield per plant.

Jing et al. (2010) study the performance of five rice genotypes derived

from different germplasm in terms of yield, harvest index (HI) and grain quality at

eight agro-ecological sites of the tropics and subtropics across Asia.

2.6.2 Heritability and Genetic Advance


Verma et al. (2000) reported high heritability coupled with high genetic

advance for plant height and days to 50% flowering.

Seetharamaiah et al. (2001) evaluated sixty-four rice genotypes for floral

and morphological traits. High heritability coupled with high-expected genetic

advance in case of panicle excertion and plant height was observed indicating

additive gene action in governing these traits.

Kavitha and Reddi (2002) studied variability and heritability in 21

genotypes of rice. Considerable amount of variability was observed for all the

quantitative characters studied. High heritability coupled with high genetic

advance was observed for number of spikelets per panicle and number of filled

grains per panicle, indicating the predominance of additive gene action in

controlling these characters.

Chandra and Pradhan (2003) studied estimates of genetic variability,

heritability and genetic advance in 65 low land rice genotypes. Phenotypic

coefficient of variation (PCV) was higher than genotypic coefficient of variation

(GCV) for all the 12 characters studied indicating the influence of environment on

the characters. A moderate to high degree of heritability estimates and genetic

advance was observed for days to 50 per cent flowering, plant height and grains

per panicle. Moderate heritability values with low genetic advances were observed

for panicle length, plot yield, 100-grain weight, grain yield per plant and harvest

index.

Sarma and Bhuyan (2004) while studying fifty eight Ahu rice genotypes in

Assam reported highest “Broad-Sense Heritability” for plant height, followed by

days to flowering and number of effective panicles per plant. Grains per panicle
and the number of effective panicles per plant showed high heritability and genetic

advance.

Choudhary et al. (2004) studied genetic variability and genetic advance for

plant traits like kernel length, panicle length, effective tiller per plant, fertile

spikelet per panicle, spikelet density, biological yield and grain yield per plant and

harvest index. All these traits were exhibiting high heritability coupled with high

genetic advance and genetic variability.

Shukla et al. (2004) carried out estimation of genetic variability,

heritability and genetic advance for 12 characters in 39 “new plant type” tropical

Japonica lines along with 3 control cultivars. The genotypes showed a wide range

of variation for all the characters. Grain yield per plant, biological yield per plant,

number of tillers and panicles per plant had high values of genetic coefficient of

variation and phenotypic coefficient of variation. Grain yield per plant, biological

yield per plant, harvest index and number of grains per panicle had high

heritability with high genetic advance.

Sinha et al. (2004) worked out estimates of variability, heritability and

genetic advance in 19 local midland landraces of rice of Surguja district along with

IR-36 for yield and its attributing characters. High genotypic coefficient of

variation was observed in grain yield followed by test weight and panicles per

plant. High heritability with high genetic advance was found for grain yield

followed by test weight and panicles per plant.

Saxena et al. (2005) evaluated forty-eight scented rice genotypes in Raipur,

Chhattisgarh, India, during the kharif season of 2001 for genetic variation in seed

yield and yield-related traits


Madhavilatha et al (2005) evaluated Fifty-four elite rice genotypes for their

variability with regards to grain yield, yield components.

Das et al. (2005) assessed 22 semi deep-water rice genotypes for genetic

variability, heritability and genetic advance. Amongst the trait under study highest

GCV was recorded for effective ear bearing tillers followed by grain yield and

plant height. Heritability in broad sense was observed to be high for all the traits

under study. The traits effective ear bearing tillers, grain yield and plant height

exhibited both high heritability and high genetic advance.

Mall et al. (2005) revealed highly significant differences among the 35

genotypes for all the characters. A wide range of variation was observed for mean

values of plant height, days to panicle initiation, days to 50 per cent flowering,

number of tillers per plant, number of panicles per plant, number of spikelets per

panicle, panicle length, flag leaf length, flag leaf width.

Singh et al. (2005) studied seven characters in 20 rice genotypes including

a local check during Kharif 2004. They reported high heritability accompanied

with high expected genetic advance for plant height indicating usefulness of

simple selection for the trait.

Satyanaryan et al. (2005) revealed high variability, heritability and genetic

advance for number of grains per panicle, spikelets fertility, days to 50% flowering

and plant height, while test weight recorded high variability and heritability

coupled with low genetic advance. The number of effective tillers per plant and

panicle length had low heritability.

Panwar (2005) studied 47 genotypes of rice including 16 local basmati

varieties and 31 high yielding varieties/advance lines to assess the genetic


variability, heritability and genetic advance for panicle characters under two

sowing dates. Highly significant variations among the genotypes were observed

for different characters. The difference between genotypic and phenotypic

coefficient of variations were relatively low for almost all the characters except

number of primary branches per panicle. High to moderate genotypic coefficient

of variation with high heritability and genetic advance were recorded for grain

yield per panicle, chaffy grain, grain yield per plant and number of filled grains.

Baber et al. (2007) reported that broad sense heritability estimates for grain

yield per plant, flag leaf width, plant height, and panicle length.

Padmaja et al. (2008) found that Heritability and genetic advance were

high for all the characters except days to 50% flowering and panicle length, which

had moderate genetic advance along with high heritability indicating the

involvement of additive type of gene action in controlling these characters.

Bisne et al. (2009) found that high heritability coupled with high genetic

advance exhibited by harvest index, total number of chaffy spikelets per panicle

grain yield per plant, total number of filled spikelet per panicle and spikelet

fertility percentage.

2.7 CORRELATIONS AND PATH-COEFFICIENT ANALYSIS

2.7.1 Correlation

Grain yield of plant is influenced by a number of components characters

and environment either directly or in directly. Contribution of each character

towards increase in yield varies from crop to crop. Correlation coefficient is

therefore used to measure the mutual relationship between various plant characters
and to determine the component characters, on which selection can be based for

genetic improvement in the yield. The correlation among the traits can be

determined in terms of phenotypic correlation and genotypic correlation.

Phenotypic correlation the association between two variables that can be

directly observed. It includes both genotypic and environmental effects. While the

genotypic or genetic correlation is the inherent or inheritable association between

two variables is. Correlation studies provide better understanding of yield

components which helps the plant breeder during selection. (Robinson et al., 1951,

Johnson et al., 1955).

Tomar et al. (2000) reported that grain yield per plant was positively

associated with plant height, number of effective tillers, panicle length, number of

grains per panicle, harvest index, biological yield and days to 50 per cent

flowering.

Nayak et al. (2001) reported that grain yield showed significant positive

correlation with plant height, panicle number per plant, panicle length, total

number of spikelets per panicle and total number of grains per panicle at both

genotypic and phenotypic levels.

Iftekharuddaula et al. (2002) studied correlation and path coefficient

analysis among nineteen genotypes of irrigated rice. Harvest index, panicle and

1000-grain weight showed significant positive correlation with grain yield.

Mishra and Verma (2002) reported that grain yield per plant was

significantly and positively correlated with number of ear bearing tillers per plant,

biological yield and harvest index at genotypic and phenotypic levels. Path

analysis indicated that spikelet density had positive direct effect on grain yield,
followed by panicle length, harvest index, biological yield, plant height. Spikelets

per panicle showed positive direct effect on harvest index followed by sterility

percentage, number of ear-bearing tillers per plant, 100-grain weight and grain

yield per plant.

Rajamani et al. (2004) reported that grain yield per plant was positively

associated with plant height, days to 50 per cent flowering, panicle length, total

number of spikelet per panicle, number of filled grains per panicle and 100 grain

weight. These characters also were positively associated among themselves.

Yogameenakshi et al. (2004) reported that number of grains per panicle

and harvest index exhibited highly positive correlation with grain yield and also a

positive intercorrelation among themselves.

Pantuwan et al. (2004) reported moderate to high heritability for grain

yield under drought has a positive indirect response to selection for grain yield

which can be achieved from selection based on drought score measured in dry

season screening.

Souroush et al. (2004) evaluated 36 genotypes of rice to determine the

relationships between grain yield and its components. The grain yield had a

positive and significant genotypic correlation with, grain weight, grain maturity

and days to 50 per cent flowering. The results of path analysis indicated that the

number of panicles per plant and number of filled grains per panicle were the most

important components with high direct effects on grain yield.

Patil and Sarawgi (2005) reported that grain yield per plant had positive

significant correlation with ear bearing tillers per plant, filled grains per panicle,

plant height and days to 50% flowering. The positive and significant correlation of
head rice recovery percentage was also observed with plant height, filled grain per

panicle, panicle length, ear bearing tillers and elongation ratio.

Anurag et al. (2006) studied path coefficients values in respect of ten

quantitative traits for 47 rice genotypes. In path coefficient analysis, panicles per

plant exhibited maximum positive direct effect on grain yield per plant .Other

characters with positive direct effects were days to 50% flowering, panicle length.

Negative direct effect was found in plants height and tillers per plant.

Girish et al. (2006) worked on F8 recombinant inbred aerobic rice

population developed by single seed descent method. They reported that grain

yield per plant was positively correlated with plant height, number of tillers,

panicle length, number of panicles plant-1, test weight, straw weight, biomass

plant-1, harvest-index and grain breadth both at genotypic and phenotypic levels.

Bisne et al. (2006) revealed the positive direct effect of characters harvest

index and 100-grain weight on grain yield. Harvest index and 100-grain weight

recorded high positive correlation with significant positive direct effect on grain

yield.

Johnson et al. (2007) studied genotypic and phenotypic correlation among

14 characters in 16 low land rice genotypes. They reported grain yield per plant

was positively correlated with plant height, biological yield and harvest index at

genotypic level. They also showed panicle length was positively correlated with

grain yield at phenotypic level.

Arumugam et al. (2008) conclude that grain yield was significantly and

positively correlated with its component characters like, 1000-grain weight and

days to 50 per cent flowering irrespective of the environment.


2.7.2 Path coefficient Analysis

The selection of a genotype is based on the study of various characters

contributing towards yield of a crop plant. The yield being quantitative trait is

dependent on the various direct and indirect added by the traits. The selection of

the traits contributing directly to the yield will be beneficial for the breeder in

changing the genetic architect of the crop plant for promoting the yield potential of

the crop. Path analysis is the effective statistical tool to determine the direct and

indirect contributions of different characters towards yield.

Path coefficient is a standardized partial regression coefficient which

separates the correlation coefficient into its components of direct and indirect

effects. Thus it helps in examining the relative contribution of both the direct and

indirect effects of independent variables on dependent variable. This technique

was originally developed by Wright (1921) and was used for the first time in

plants by Dewey and Lu (1959).

Chakraborty et al. (2001) revealed high positive direct effects of harvest

index, panicle length and 100-grain weight on the grain yield per plant, indicating

the possibility of yield improvement through direct selection for these traits.

Mishra and Verma (2002) reported that grain yield per plant was

significantly and positively correlated with number of ear bearing tillers per plant,

biological yield and harvest index at genotypic and phenotypic levels. Path

analysis indicated that spikelet density had positive direct effect on grain yield,

followed by panicle length, harvest index, biological yield, plant height. Spikelets

per panicle showed positive direct effect on harvest index followed by sterility
percentage, number of ear-bearing tillers per plant, 100-grain weight and grain

yield per plant.

Satish et al. (2003) worked out genotypic and phenotypic correlations and

path coefficient analysis for 13 quantitative and physiological characters

influencing the yield in rice. Number of productive tillers per hill and number of

grains per panicle exhibited significant positive correlation with yield. Path

coefficient analysis revealed that number of productive tillers per hill and number

of grains per panicle showed higher positive direct effect on yield.

Murthy et al. (2004) revealed a significant and positive correlation of grain

yield per plant with days to flowering, days to maturity and leaf length.

Khedikar et al. (2004) revealed that the path coefficient analysis of test

weight had the highest positive direct effect on grain yield per plant. It was

followed by spikelet density, effective tillers per plant, panicle length and days to

50% flowering, and hence direct selection through these characters is more

effective.

Souroush et al. (2004) evaluated 36 genotypes of rice to determine the

relationships between grain yield and its components. The grain yield had a

positive and significant genotypic correlation with, grain weight, grain maturity

and days to 50 per cent flowering. The results of path analysis indicated that the

number of panicles per plant and number of filled grains per panicle were the most

important components with high direct effects on grain yield.

Naik et al. (2005) reported that plant height had the positive and significant

correlation with panicle length and biological yield per plant at phenotypic and

genotypic level. Path coefficient analysis of different characters contributing


towards grain yield per plant revealed that paddy length had the highest positive

direct relationship with grain yield per plant followed by paddy breadth, biological

yield per plant and effective tillers per plant.

Bisne et al. (2006) revealed the positive direct effect of characters number

of spikelets per panicle, harvest index and 100-grain weight on grain yield.

Harvest index and 100-grain weight recorded high positive correlation with

significant positive direct effect on grain yield.

Manonmani and Ranganathan (2006) reported that number of productive

tillers plant-1 had high positive direct effect on yield followed by days to 50 per

cent flowering, plant height, grains panicle-1 and 100-grain weight. Days to 50 per

cent flowering had positive indirect effect on yield through plant height. Number

of grains panicle-1 also contributed indirect positive effect on yield through days to

flowering, plant height and number of productive tillers. 100-grain weight

contributed equally through direct and indirect effects.

Anurag et al. (2006) studied path coefficients values in respect of ten

quantitative traits for 47 rice genotypes. In path coefficient analysis, panicles per

plant exhibited maximum positive direct effect on grain yield per plant .Other

characters with positive direct effects were days to 50% flowering, panicle length.

Negative direct effect was found in plants height and tillers per plant.

Panwar and Ali (2007) reported that grain yield per panicle had the highest

direct effect on grain yield per plant followed by harvest index; biological yield

per plant and productive tillers per plant suggesting that the improvement in grain

yield could be efficient, if the selection is based on these component characters.


CHAPTER III

MATERIALS AND METHODS

The present investigation “Effect of Drought Tolerant QTL for Yield

and Yield Parameters under Managed Water Stress as Well as Non Stress

Condition” was conducted during Kharif 2010 at Instructional cum Research

Farm of, Indira Gandhi Krishi Vishwavidyalaya, Raipur, Chhattisgarh, India.

Location and climate:

Raipur is situated in the central part of Chhattisgarh at 21016‟‟ N latitude

and 81036‟‟ E longitude with an altitude of 289.56 meters above mean sea level.

The climatic condition of Raipur is hot and sub-humid type.

The details of materials used and the methodologies applied in the

investigation are described below:

Experimental details:

The experimental material for the present investigation comprised 77 lines

derived from Nagina 22/ Swarna cross including checks (MTU-1010, IR-36, N-22,

Swarna & IR-64) the experimental material was received from IRRI through

CRRI, Cuttack. The details of these lines are presented in table 3.1.
Appendix II: List of genotype

EN0 DESIGNATION QTL information EN0 DESIGNATION QTL information


1 IR 86931-B-6 with QTL DTY3.2 39 IR 86931-B-323 with QTL DTY1.1
2 IR 86931-B-9 with QTL DTY3.2 40 IR 86931-B-326 without any QTL
3 IR 86931-B-26 with QTL DTY1.1 41 IR 86931-B-310 with QTL DTY1.1
& 3.2
4 IR 86931-B-32 with QTL DTY3.2 42 IR 86931-B-346 with QTL DTY1.1
5 IR 86931-B-36 without any QTL 43 IR 86931-B-347 with QTL DTY1.1 & 3.2
6 IR 86931-B-40 without any QTL 44 IR 86931-B-365 with QTL DTY3.2
7 IR 86931-B-50 without any QTL 45 IR 86931-B-366 with QTL DTY3.2
8 IR 86931-B-55 with QTL DTY1.1 46 IR 86931-B-373 with QTL DTY1.1 & 3.2
& 3.2
9 IR 86931-B-56 with QTL DTY1.1 47 IR 86931-B-390 with QTL DTY3.2
& 3.2
10 IR 86931-B-60 with QTL DTY3.2 48 IR 86931-B-400 with QTL DTY1.1
11 IR 86931-B-66 with QTL DTY1.1 49 IR 86931-B-414 with QTL DTY1.1 & 3.2
12 IR 86931-B-67 with QTL DTY3.2 50 IR 86931-B-431 without any QTL
13 IR 86931-B-78 without any QTL 51 IR 86931-B-454 with QTL DTY3.2
14 IR 86931-B-86 with QTL DTY1.1 52 IR 86931-B-456 with QTL DTY1.1 & 3.2
15 IR 86931-B-103 without any QTL 53 IR 86931-B-464 with QTL DTY3.2
16 IR 86931-B-104 without any QTL 54 IR 86931-B-467 with QTL DTY3.2
17 IR 86931-B-120 without any QTL 55 IR 86931-B-486 with QTL DTY3.2
18 IR 86931-B-131 with QTL DTY1.1 56 IR 86931-B-487 without any QTL
19 IR 86931-B-132 with QTL DTY3.2 57 IR 86931-B-498 with QTL DTY1.1
20 IR 86931-B-147 with QTL DTY1.1 58 IR 86931-B-502 with QTL DTY1.1 & 3.2
21 IR 86931-B-150 with QTL DTY1.1 59 IR 86931-B-516 with QTL DTY1.1 & 3.2
22 IR 86931-B-152 without any QTL 60 IR 86931-B-519 with QTL DTY1.1 & 3.2
23 IR 86931-B-158 without any QTL 61 IR 86931-B-17 with QTL DTY1.1
24 IR 86931-B-142 without any QTL 62 IR 86931-B-521 with QTL DTY1.1
25 IR 86931-B-166 with QTL DTY1.1 63 IR 86931-B-523 with QTL DTY3.2
& 3.2
26 IR 86931-B-170 with QTL DTY3.2 64 IR 86931-B-524 with QTL DTY1.1 & 3.2
27 IR 86931-B-173 without any QTL 65 IR 86931-B-547 with QTL DTY3.2
28 IR 86931-B-187 with QTL DTY1.1 66 IR 86931-B-549 with QTL DTY3.2
29 IR 86931-B-192 with QTL DTY1.1 67 IR 86931-B-552 with QTL DTY1.1 & 3.2
30 IR 86931-B-194 without any QTL 68 IR 86931-B-559 without any QTL
31 IR 86931-B-224 with QTL DTY1.1 69 IR 86931-B-561 with QTL DTY3.2
32 IR 86931-B-233 with QTL DTY1.1 70 IR 86931-B-564 with QTL DTY1.1
33 IR 86931-B-256 with QTL DTY1.1 71 IR 86931-B-574 with QTL DTY3.2
& 3.2
34 IR 86931-B-263 without any QTL 72 IR 86931-B-578 without any QTL
35 IR 86931-B-279 with QTL DTY1.1 73 N22 Check
& 3.2
36 IR 86931-B-294 with QTL DTY1.1 74 Swarna Check
& 3.2
37 IR 86931-B-301 with QTL DTY1.1 75 IR 64 Check
38 IR 86931-B-311 without any QTL 76 IR 36 Check
77 MTU 1010 Check

3.2 Methods

3.2.1 Field Studies

The lines derived from a cross between Nagina 22/ Swarna rice cultivar

were evaluated in the field during wet season 2010 at research farm of IGKV,

Raipur. The field trials were conducted under terminal stage drought (TSD) and

irrigated control conditions. The fields selected for the study were upland in

topology with good drainage and percolation rate and had sandy loam soil. Seeds

were sown in nursery on 15/07/2010 and 01/07/2010 for TSD and irrigated

conditions respectively. Under TSD each genotype was sown in two rows of 3.4

m. length, while under irrigated condition two rows of 2 m. length with spacing of

20 cm between rows. All the genotypes were replicated twice in RCBD design

(Plate I). All normal packages of practices were followed to raise a good crop. The

total rainfall during crop cycle wet season 2010 was 1065 mm. 86% of the rainfall

was received during vegetative phase and 13% rainfall at reproductive phase

which are sown in fig. 3.1.


Fig 3.1: Distribution of rainfall during wet season 2010

Observations:

Observation recorded under field conditions are described below:-

Days to 50% flowering:-

The time taken (days) from sowing to flowering period was recorded

on plot basis by visual means.

Plant height:-

Plant height was measured in centimeter (cm) from soil surface to tip of

the tallest panicle at maturity.

Panicle length:-

Single panicle length of 3 randomly selected plants was measured in

centimeters from base to tip of panicle and was averaged.

Flag leaf length:-

Flag leaf length of 3 randomly selected plants was recorded by

measuring the length (cm) from nod to tip of flag leaf and was averaged.

Flag leaf width:-


Flag leaf width (cm) was recorded by measuring the leaf width at the

central part of the leaf.

Flag leaf Area

It is the product of Flag leaf Length and Flag leaf Width

Grain length

The length of randomly selected twenty spikelets was measured in

terms of millimeters.

Grain Breadth

The breadth of randomly selected twenty spikelets was measured in

terms of millimeters.

Grain L: B ratio

The length/breadth ratio of randomly selected ten spikelets was

calculated by dividing respective length with breadth.

100 seed weight:-

Test weigh was recorded for 100 grains of each line in grams.

Harvest index:-

Harvest index was worked out by using the formula as given below:

Harvest index = (Grain yield/ Biological yield) *100

Biological yield:-

The biological yield was recorded as actual weight in gram of total

biomass of shoot portion per plot and later converted to g/ m2 for further

analysis.
Grain yield:-

The actual yield of grain in gram per line was recorded and later

converted to g/ m2 for further analysis.

Statistical analysis:-

The data recorded on all the traits related to yield and yield

contributing characters in both the season were statistically analyzed for

variance.

Analysis of variance:-

The mean data of each replication was used for analysis of variance using

RCBD design. The mean data were subjected to variance analysis and test of

significance as per the method of Fisher (1935).

Parameters of variation:

Mean

Mean is the average value of observation of genotypes of a series. It

represents the standard average value over fluctuation in the environment. Mean

was calculated by the following formula:

X = ∑Xi / n

Where,

∑Xi = Summation of all the observations

n = Total number of observations

Range:-
Range is the difference between the highest and the lowest value of a series

of observations and thus provides the information about the extent of variability

present in the genotypes.

Range = Highest value - Lowest value

Genetic Analysis:-

Variability:-

Genotypic coefficient of variation (GCV) and phenotypic coefficient of

variation (PCV) were calculated by the method suggested by Burton (1952). The

estimates of PCV and GCV were classified as low, moderate and high according to

Sivasubramanian and Madhavamenon (1973).

< 10 per cent = Low

10 – 20 per cent = Moderate

> 20 per cent = High

Heritability (broad sense):-

It is the ratio of genotypic variance to the phenotypic variance (total

variance). Heritability for the present study was calculated in a broad sense by

adopting the formula as suggested by Hanson et al. (1956). The estimates of

heritability broad sense were classified as low, moderate and high according to

Lafitte et al. (2004).

0.2 – Low

0.6 – Moderate

0.9 – High

Genetic advance:-
Improvement in the mean genotypic value of selected plants over the

parental population is known as genetic advance. Expected genetic advance (GA)

was calculated by the method suggested by Johnson et al. (1955)

Genetic advance as percentage of mean:-

It was calculated by the following formula:

GA as percentage of mean = (Genetic Advance/ General mean) x 100

GA was categorized as:

Range Category

< 10 per cent Low

10 - 20 per cent Moderate

> 20 per cent High

Correlation Coefficient:-

Correlation coefficient (r) was calculated for grain yield and yield

contributing characters by using the standard procedure given by Searle (1961).

Path analysis

Path analysis was originally developed by Wright (1921) and first used for

plant selection by Dewey and Lu (1959). It measures the direct and indirect

contribution of independent variables on dependent variable.


The results of path coefficient analysis are interpreted as per the following

scale suggested by Lenka and Mishra (1973).

Value of direct and indirect effects Rate/ Scale

0.00 to 0.09 Negligible

0.10 to 0.19 Low

0.20 to 0.29 Moderate

0.30 to 0.99 High

>1.00 Very High


CHAPTER IV

RESULTS AND DISCUSSION

The present investigation entitled “Effect of drought tolerant QTLs on

yield and yield contributing traits under managed water stress as well as non

-stress conditions” was carried out with the main objectives of genetic analysis

and to understand the effect of drought tolerant QTLs on yield under stress as

well as non stress conditions. The 77 lines with and without QTLs along with

checks were phenotype under two set of environmental conditions, one under

irrigated condition and another under terminal stage drought condition during wet

season 2010. The mean data of irrigated and TSD condition were used for genetic

analysis. The results, thus obtained are presented under following headings:

4.1 General observations

4.2 Variability

4.2.1 Analysis of variance

4.2.2 Drought Susceptibility Index

4.2.3 Mean and variability parameters

4.2.4 Genotypic and phenotypic coefficient of variation

4.2.5 Heritability and genetic advance

4.3. Correlation Studies

4.4 Path Analysis

4.5 Effect of QTL for Yield and yield contributing traits

4.1 General observations


Overall, during the crop season total rainfall of 1065 mm. was received,

The erratic rainfall along with management of field (including selection of fields

with light textured soil high in topology, good drainage and agronomic

adjustment) has resulted in substantial imposition of stress. The drought intensity

index for this trial during wet season was 0.732 which indicate imposition of very

high level of water stress. In most breeding programme screening for drought

tolerances fail to impose a sufficiently sever stress in their trials and this are not

able to accurately predict drought tolerant line (Kumar et al. 2008)

4.2 Variability

4.2.1 Analysis of variance

The experimental material for the present investigation comprised 77 lines

derived from Nagina 22/ Swarna cross including checks (MTU-1010, IR-36, N-22,

Swarna & IR-64) all the genotypes were evaluated in wet season 2010 for various

phonological, agronomical and physiological traits. The data on various characters

were recorded either on five plants basis or on plot basis. The mean of five plants

or plot data of each replication was subjected to Randomized Complete Block

Design (RCBD) analysis. The results of RCBD analysis under Irrigated and TSD

condition of wet season 2010 is presented in table 4.1. Significant variation was

noticed among the 77 lines for grain yield and yield contributing traits under

rainfed as well as TSD situation. These results were in agreement with the results

of Singh et al. (1984) and Monalisa et al. (2006) as they have also reported lots of

genetic variability for these traits.

4.2.2 Drought Susceptibility Index (DSI)


The mean data of grain yield of individual lines for both conditions

separately and Drought Susceptibility Index of the population were statistically

analyzed. The results of this analysis are presented in table 4.2.

A perusal of Drought Susceptibility Index data indicated highest value for

check genotype Swarna. this variety had grain yield of 211.1 g/m2 under irrigated

condition while it produced 10.0 g/m2 under stress condition exhibiting severe

reduction in yield which is indicated by high DSI value. This genotype is known to

be susceptible.

The lower value for Drought susceptibility index was observed for IR

86931-B-347 (-0.07) and IR 86931-B-456 (-0.07) which indicates low or virtually

no reduction in yield under stress condition compared to irrigated condition.

However, these two genotype also yielded substantially low under irrigated

condition as well, therefore the drought susceptibility index value for these

genotype is mis-leading, however, these were number of lines with grain yield

more than overall average under irrigated condition and low drought susceptibility

index indicating tolerance to water stress.

4.2.3 Mean and variability parameters

The mean data of individual lines of the population were statistically

analyzed to generate overall mean and other parameters. The results of this

analysis are presented in table 4.3.

Grain yield under irrigated condition ranged from 122.20 to 1022.20 g/ m2

with population mean of 432.02 g/ m2, while under TSD condition it ranges from

10.00 to 173.35 g/ m2 with a mean of 115.53 g/ m2 which indicates marked

differences among the 77 genotype population for grain yield under stress
condition, which indirectly indicates the lines have ability to cope with water

stress. The mean reduction in grain yield of population due to TSD over Irrigated

condition was 81.35%, indicating very high stress levels. Such high stress levels

were desirable because a high percentage reduction of yield is necessary to remove

the effect of yield potential and clearly identify lines that are drought resistance

(Babu et al., 2003; Lafitte et al., 2006). Similar results of reduction in yield were

reported by Lanceras et al. (2004).

Days to 50% flowering of 77 genotypes under irrigated condition ranged

from 71 to 97 days with mean of 83 days while under TSD condition it ranged

from 67 to 92 days with mean of 78 days. The standard deviations were

comparable over the season. Under same temperature condition it was reported to

increased days to 50% flowering under drought condition .While in this

experiment flowering was not delayed, it was mainly due to delayed planting

(most of the rice genotypes are at least partially photo and thermo sensitive) and

water stress was imposed at the time of reproductive phase. Usually vegetative

stage drought result is delay in flowering

Biomass production (biological yield) of 77 genotype population under

irrigated condition ranged from 789 to 3322 g/ m2 with mean of 1928.85 g/ m2,

while under TSD condition it ranged from 133 to 566.65 g/ m2 with mean of

416.75 g/ m2. The mean and range is higher in irrigated compare to TSD

Condition.

There was marked difference in population means for plant height of TSD

and Irrigated. Plant height of 77 lines under irrigated condition ranged from 79.65
cm to 176 cm with mean of 144.83 cm, while under TSD condition it ranged from

75.65cm to 150.80 cm with mean of 123.90 cm.

100 Grains weight of 77 lines under irrigated condition ranged from 1.62 g

to 2.60 g with mean of 2.17 g, while under TSD condition it ranged from 1.49 g to

2.54 g with mean of 2.07 this was mainly because of shriveled grain production

under stress.

Harvest index (HI) of 77 lines under irrigated condition ranged from 10.98

to 43.32 with mean of 22.12, while under TSD condition it ranged from 3.00 to

39.20 with mean of 27.82

Panicle length of 77 lines under irrigated condition ranged from 20.50 cm

to 29.20 cm with mean of 25.31 cm, while under TSD condition it ranged from

17.20 cm to 28.85cm with mean of 23.81 cm.

Flag leaf length of 77 lines under irrigated condition ranged from 27.05

cm to 49.00 cm with mean of 36.59 cm, while under TSD it ranged from 28.05 cm

to 55.00 cm with mean of 41.61 cm.

Flag leaf width of 77 lines under irrigated condition ranged from 1.10 cm

to 1.70 cm with mean of 1.37 cm, while under TSD condition it ranged from 0.85

cm to 1.65cm with mean of 1.23 cm.

Flag leaf length: width ratio of 77 lines under Irrigated condition ranged

from 25.70cm to 45.25cm with mean of 35.12cm. While under TSD it ranged from

23.35cm to 51.45cm with mean of 35.70cm.


Grain length of 77 lines under irrigated condition ranged from 7.43mm. to

10.03 mm. with mean of 83.34 mm, while under TSD condition it ranged from

6.68 mm to 9.93 mm with mean of 8.14 mm.

Grain breadth of 77 lines under irrigated condition ranged from 1.95 mm to

3.15 mm. with mean of 2.69 mm, while under TSD condition it ranged from 2.05

mm. to 3.10 mm. with mean of 2.73 mm.

Grain L: B ratio of 77 lines under Irrigated condition ranged 2.12 mm to

4.01mm with mean of 3.12 mm. while under TSD it ranged 2.51 mm to 3.77 mm

with mean of 3.00 mm.

Flag leaf length, flag leaf width, grain length, grain width, grain L: B ratio,

panicle length and 100 grain weight of population means did not exhibited much

difference across the moistures regime.

4.2.4 Genotypic and phenotypic coefficient of variation

To develop a high yielding genotypes coupled with drought tolerance,

populations showing high variability serves prime source for effective selection. In

the present investigation high phenotypic and genotypic coefficient of variation

was obtained for most of the traits (Table 4.3).

Panicle length, 50% flowering, grain length, grain breadth and Grain L: B

ratio showed low GCV and PCV under both Irrigated and TSD condition. Low

value of GCV for Grain width, Grain length and Grain L: B ratio is not similar to

Husain et al. (1987).


Plant height and flag leaf length, showed moderate GCV and PCV under

both Irrigated and TSD condition. Similar result of GCV and PCV is also reported

by Umadevi et al. (2009).

Grain yield, showed high GCV and PCV under both Irrigated and TSD

condition. Biological yield, show moderate GCV and high PCV under TSD

condition and high GCV and PCV under Irrigated condition. While harvest index

show high GCV and PCV under both condition.

GCV and PCV for grain yield under irrigated condition were 39.31% and

46.94 % while under TSD condition it was 25.77 % and 29.49 %, respectively.

GCV and PCV values under individual set of environmental condition indicate that

influence of environment on the expression of character is low. Due to high GCV

and PCV selection of character will be rewarded.

Days to 50% flowering showed fairly consistent but low estimates of GCV

and PCV, 5.61 and 5.63 respectively under Irrigated condition and 5.25 and 5.42

respectively under TSD condition, across the moisture regime. Similar finding

were also reported by Singh et al. (1999) and Monalisa et al. (2006).

Biological yield showed fairly higher values of GCV and PCV 24.61 and

30.99 respectively under Irrigated condition and 16.83 and 19.58 respectively

under TSD condition. Comparable GCV and PCV were observed for harvest index

24.61 and 31.44 respectively under Irrigated and 19.11 and 23.79 respectively

under TSD condition, across the environmental conditions. Similar estimates of

variations for most of the traits were also reported by Singh et al. (1999).
All the characters showed little difference between PCV and GCV which

indicate that traits were lesser influence by environment. All the characters showed

high PCV than GCV and it was also reported by Umadevi et al. (2009).

4.2.5 Heritability and genetic advance

Heritability and genetic advance are important selection parameter.

Heritability estimates along with genetic advance are more useful in predicting the

gain under selection then the heritability alone. Table 4.3 shows the heritability

and genetic advance of yield and yield contributing traits. When heritability and

genetic advance are high than selection is effective due to additive gene effect and

when both are low than selection is not effective due to non additive gene effect.

Grain yield under irrigated condition showed medium heritability 0.70

coupled with high genetic advance as percentage of mean 67.82, while under TSD

condition high heritability was 0.76 with genetic advance as percentage of mean

was 46.39. Similar results of medium heritability for Grain yield was observed in

numerous studies under drought condition (Babu et al., 2003; Yue et al., 2005;

Venuprasad et al., 2007, 2008; Kumar et al., 2008; Bernier et al., 2009a; Verulkar

et al., 2010). Moderate to high heritability for Grain yield under drought stress

indicated selection for yield under drought stress is repeatable and support direct

selection for yield rather than secondary traits related to drought tolerance.

Further highest genetic advance as percentage of mean indicated the

presence of additive gene action and selection will be rewarding for Grain yield.

Biological yield and Harvest index showed moderate heritability 0.63, and 0.61

respectively under Irrigated condition and 0.73 and 0. 64 respectively under TSD

condition accompanied with higher genetic advance as percentage of mean 40.25


and 39.69 respectively under irrigated condition and, 29.78 and 31.63 respectively

under TSD condition, indicating selection of these characters can also be used as

criteria for selection of genotypes with higher yield. A similar result of

comparatively lower heritability under Irrigated condition was reported to observe

by Babu et al. (2003).

Heritability of 0.63 and 0.73 coupled with 40.25 and 29.78 genetic

advances as percentage of mean was observed for Biological yield under Irrigated

and TSD condition respectively. This result also showed that Biological yield can

be included in the criteria of selection for Grain yield under drought condition.

Similar estimates of high heritability coupled with genetic advance were reported

by Gomez and Kalamani (2003) and Bernier et al. (2007).

Days to 50% flowering and plant height showed higher heritability 0.99

and 0.72, respectively under irrigated condition and 0.93 and 0.83, respectively

under TSD condition with medium genetic advance as percentage of mean, 11.51

and 25.20, respectively under irrigated and 10.51 and 23.49, respectively under

TSD condition which indicate that selection based on these characters will not be

rewarding due to non additive gene action. The high heritability is being exhibited

due to favorable influence of environment rather than genotype and selection for

such trait may not be rewarded. Similar results were found by Umadevi et al.

(2009).

Plant height showed high heritability (0.83), accompanied with high

genetic advance as percentage of mean (23.47) under TSD condition. Heritability


and genetic advance as percentage of mean was not comparable under Irrigated

condition (0.72 and 25.20 respectively) for Plant height.

Days to 50% flowering showed high heritability (0.99) coupled with

moderate GA as percentage of mean (11.51) under Irrigated condition, while

under TSD condition it showed 0.93 h2 and 10.51 genetic advances as percentage

of mean, indicating selection based on higher days to 50% flowering is not

rewarding. A similar finding of result was revealed by Gomez and Kalamani

(2003). Higher heritability for 50% flowering and Plant height is also reported by

Verulkar et al. (2010).

High heritability (0.99, 0.72and 0.63) and moderate genetic advance as

percent of mean (11.51, 25.20 and 40.25) showed for days 50% flowering was

recorded in Plant height and biological yield respectively under Irrigated

condition and these traits showed also high values of heritability and moderate

genetic advance as percentage of mean under TSD condition.

4.3 Correlation Studies

Grain yield of plant is not unitary character but is influenced by a number

of components characters and environment either directly or in directly.

Contribution of each character towards increase in yield varies from crop to crop.

Correlation coefficient is therefore used to measure the mutual relationship

between various plant characters and to determine the component characters, on

which selection can be based for genetic improvement in the yield.


The mean data for each character under two different sets of condition

were first classified into group depending on presence or absence of different DT

QTLs within group the data was analyzed for correlation among different traits.

4.3.1 Correlation among variable in QTL DTY 1.1 lines

The character Days to 50% flowering showed positive correlation under

TSD condition and negative correlation with yield under irrigated condition (Table

4.4).

Plant height showed negative and significant correlation with Harvest

index under TSD and irrigated condition respectively and with biological yield

they showed negative correlation under TSD & irrigated condition.

Panicle length showed negative and significant correlation with harvest

index and with grain yield it show positive correlation under irrigated condition

Flag leaf length showed positive & highly significant and significant

correlation with Flag leaf area under irrigated and TSD condition respectively and

it also show positive and significantly correlated with biological yield under

irrigated condition it showed negative correlation in TSD and positive correlation

in irrigated condition with grain yield

Highly significant positive correlation of Flag leaf width was observed

with Flag leaf area under TSD condition while in irrigated condition it showed

positively significant correlation. Flag leaf area showed positive correlation with

grain yield under both condition but non significant.


The attribute biological yield showed positive and highly significant

correlation with grain yield under irrigated while in TSD condition they showed

positive correlation but non significant

Harvest index is showing highly significant and positive correlation with

Grain yield under both TSD & irrigated condition means grain yield increase % of

Harvest index.

Grain length is showing positive and highly significant correlation with

length width ratio of grain under irrigated condition.

4.3.2 Correlation among variable in DTY 1.1 & 3.2 lines

Under correlation coefficient in QTL DTY 1.1&3.2 lines some attributes

showed positive and significant correlation and some attribute showed significant

but negative correlation with other different traits (Table 4.5).

Days to 50% showed positive correlation with grain yield under both

condition but non significant

Plant height showed positive and significant correlation with flag leaf

length but negative & significant correlation with Grain Length and length-width

ratio of grain under TSD condition. While in irrigated condition they express

negative significant correlation with harvest index. There for falling both condition

viz. quality in term of Grain length quantity in terms of Harvest index one should

prefer dwarf plant Days to 50% flowering and panicle length showed non

significant correlate under the both condition with growth yield revealing that long

pedicure are produced in late duration variety which enhances yield. Flag leaf

length showed positive & significant correlation with Flag leaf area under the both

TSD & Irrigated condition. And FLW showed positive significant correlation
under TSD condition and positively high correlated under irrigated condition with

flag leaf area

Biological yield expressed highly significant and positive correlation with

grain yield under the both TSD and irrigated condition, this reveal that irrespective

of condition the Biological yield enhances by along with it grain yield showed

positive highly significant correlation in TSD condition and positive significant

correlation in irrigated condition with character harvest index.

Harvest index showed positive significant correlation with seed index

under the irrigated condition and Grain length showed highly significant positive

correlation with length width ratio of grain under the irrigated condition only.

Grain width showed negative significant correlation with grain L: B ratio

of under the both condition. In QTL DTY 1.1&3.2 lines maximum character

4.3.3 Correlation among variable in with QTL DTY 3.2 lines

Under the correlation coefficient study among QTL DTY 3.2 lines it was

found that total eleven attributes were significantly positively correlated while

other attributes including to negative traits viz. namely Days to 50% flowering in

TSD condition (Table 4.6).

Plant height showed positive significant correlated with biological yield

under TSD condition and it also express positive correlation with grain yield under

TSD condition.

This had shown that increase in the height of the plant is associated with

increase in biological yield & grain yield.

Flag leaf length expressed highly significant positive correlation with Flag

leaf area under both TSD and irrigated condition.


Significant and positive correlation observe of Flag leaf area with Flag leaf

width under irrigated and TSD condition Flag leaf area being dependent character

shows such relationship with Flag leaf length& Flag leaf width.

Harvest index is showing highly significant and positive correlations with

biological yield under both irrigated and TSD condition

Biological yield is showing highly significant and positive correlations

with grain yield under both irrigated and TSD condition. Grain yield expressed

highly significant and positive correlation with harvest index under the both

irrigated and TSD condition.

This indicates that increase in the biological yield result in increasing the

grain yield which ultimately enhances Harvest index.

.Grain length showed highly significant and positive correlation with

length width ratio of grain under both condition while with 100 seed weight it

showed highly significant and positive correlation under TSD condition. This

reveals that increase in the Grain length increases the Seed index. Grain breadth is

showing positive & significant with 100 seed weight under TSD condition in QTL

DTY 3.2 lined maximum significant and positive relation is been seen under TSD

condition.

4.3.4 Correlation among variables in without QTL lines

Plant height is showing positive & significant correlation with panicle

length and flag leaf length under irrigated condition and observed positive

correlation with grain yield under both TSD & irrigated condition but non

significant (Table 4.7).


Highly significant and positive correlation with Flag leaf length is observed

with Flag leaf area under both TSD and irrigated condition.

Flag leaf area recorded highly significantly and positive correlation with

Flag leaf width under both TSD and irrigated condition.

Attributes Biological yield is showing highly significant and positive

correlation with grain yield under the both TSD & irrigated condition.

Grain yield expressed positive and highly significant correlation with

harvest index under irrigated condition while positive and significant correlation in

TSD condition.

Grain length showed highly significance and positive correlation with 100

seed weight under the both TSD and irrigated condition while it showed highly

significant and positive correlation with length width ratio of grain under TSD

condition.

Grain width is showing highly significant but negative correlated with L:B

ratio of grain under the both condition maximum character showed negative

correlated with grain yield under the only TSD condition

4.3.5 Correlation among variable in check lines

In correlation study under the check lines show less number of significant

and positive correlation and few characters show negative significant correlation

(Table 4.8).

Days to 50 % flowering is showing significant negative correlation with

Grain yield and harvest index correlation with while in irrigated condition it

showed significant negative correlation with plant height. Panicle length showed

negative but significant correlation with grain yield under TSD condition.
Harvest index expressed positive significant correlation with Grain yield under

irrigated and TSD condition.

Most of the character showed positive correlation with grain yield under

the both condition but non-significant.

The result of correlation coefficient among different traits (for all lines

carrying DT QTL 1.1) is presented in tables .A perusal of this table indicate

highly positive and significant correlation between grain yield with Harvest index

and biological yield under both irrigated as well as terminal stage drought

condition these result are in agreement to the finding of Tomar et al. (2000),

Iftekharuddaula et al. (2002), Mishra and Verma (2002), Bisne et al. (2006),

Johnson et al. (2007) for the character Harvest Index for the all the clusters under

both conditions; Tomar et al. (2000), Mishra and Verma (2002) and Johnson et al.

(2007) for the character Biological yield for all the clusters under both conditions.

These results clearly indicate that for both the condition increase in grain yield can

be achieved if selection is made for biological yield and harvest index. Almost

similar result were obtained for correlation among trait carrying different DT

QTLs which to same extent indicate that trait correlation are not significantly

affected by presence or absence of different DT QTLs.

The result obtained for correlation with yield is in accordance with Nayak

et al. (2001), Rajmani et al.(2004), Patil and Sarawagi (2005), Johnson et al.

(2007) for character Plant height for the cluster without QTL and Check lines

under both condition, cluster QTL DTY 3.2 under TSD condition; Tomar et al.

(2000), Rajmani et al. (2004), Souroush et al. (2004) and Patil and Sarawagi

(2005) for the character Days to 50 % flowering for the cluster 1.1 and 3.2 under
both condition for cluster QTL DTY 1.1 and QTL DTY 3.2 under TSD and

Irrigated condition respectively; Tomar et al. (2000), Nayak et al. (2001), Rajmani

et al. (2004) and Johnson et al. (20007) for the character Panicle length for the

cluster QTL DTY 3.2, QTL DTY 1.1 and 3.2 under both condition; QTL DTY 1.1,

Without QTL lines and Checks under Irrigated condition; Iftekharuddaula et al.

(2002), Rajmani et al. (2004), Souroush et al. (2004)and Bisne et al. (2006) for the

character Seed Index for all the clusters under both conditions.

4.4 Path analysis

The path analysis in the study was undertaken mainly to understand the

change in direct and indirect effect of different traits by the presence/absence of

different DT QTLs.

The correlation coefficient of yield with important field and quality

characters were partitioned into direct and indirect effects taking yield per plant as

dependent variable. The selection of a genotype is based on the study of various

characters contributing towards yield of a crop plant. The yield being quantitative

trait is dependent on the various direct and indirect added by the traits. The

selection of the traits contributing directly to the yield will be beneficial for the

breeder in changing the genetic architect of the crop plant for promoting the yield

potential of the crop. Path analysis is the effective statistical tool to determine the

direct and indirect contributions of different characters towards yield. Path


coefficient is a standardized partial regression coefficient which separates the

correlation coefficient into its components of direct and indirect effects. Thus it

helps in examining the relative contribution of both the direct and indirect effects

of independent variables on dependent variable. This technique was originally

developed by Wright (1921) and was used for the first time in plants by Dewey

and Lu (1959).

4.4.1 Path analysis in QTL DTY 1.1 lines

The correlation coefficient of various characters for lines QTL DTY 1.1

was spliced into direct and indirect effects for environment (Table 4.9).

Among the characters, harvest index (1.254,0.811) followed biological

yield (0.844, 0.701) and panicle length(0.035, 0.033) were found to exert positive

direct effect while days to 50% flowering (-0.021, -0.032), plant height(-0.011, -

0.026) were found to negative direct effect and grain length (0.174, -0.338), flag

leaf length(0.211, -0.411), flag leaf width (0.230, -0.433) and grain breadth (-

0.273, 0.251), Grain L:B ratio (-0.395, 0.356), Flag leaf area (-0.274, 0.493) and

Seed index (-0.040, 0.022) showed vice-versa effect for both condition.

The breeder should go for direct selection of the plant height (-0.011, -

0.026), flag leaf width (0.230, -0.433), grain breadth (-0.273, 0.251), harvest index

(1.254, 0.811) and biological yield (0.844, 0.701) as it exhibited direct positive

effect with yield under both conditions.

Beside these the breeder can also go for selection of character days to 50%

flowering (-0.021, -0.032), panicle length (0.035, 0.033) flag leaf area (-0.274,

0.493) grain length width ratio (-0.395, 0.356) and seed index under irrigated

condition.
On the contrary, direct effect and correlation under one condition shows

vice-versa relationship for other condition i.e. days to 50% flowering (-0.021, -

0.032), panicle length (0.035, 0.033), flag leaf area (-0.274, 0.493) grain length

breadth ratio (-0.395, 0.356) and seed index (-0.040, 0.022) for TSD condition.

Thus, in such cases, one showed consider only those indirect affecting characters

which enhance the value of correlation coefficient which ultimately enhances

yield.

4.4.2 Path analysis in QTL DTY 1.1 & 3.2 lines

Among the characters Biological yield (0.506, 0.887), Harvest index

(0.753, 0.604) and flag leaf length (0.089, 0.091) were found to exert positive

direct effect and Days to 50% flowering (-0.001, -0.051) plant height (-0.056, -

0.044), panicle length (-0.018, -0.049) were found to import negative direct effect

and flag leaf width (0.114, -0.008), grain width (0.315, -0.791) grain L:B ratio

(0.372, -1.731) seed index (0.001,-0.020) and flag leaf area (-0.124,0.012) & grain

length –(0.383, 1.451) showed vice-versa effect, under both conditions (Table

4.10).

Among them breeder should go for direct selection of the traits biological

yield (0.506, 0.887), harvest index (0.753, 0.604), plant height, (-0.056, -0.044)

and Flag leaf length (0.089, 0.091) as the kind of direct effects exerted by them is

also seen in there correlation coefficient with yield for both condition.

Beside those the breeder can also go for direct selection of characters like

flag leaf width (0.114, -0.008), grain weight (0.315,-0.791) and seed index (0.001,

-0.020) keeping in view of their association with yield. under TSD condition while

grain length (-0.383, 1.45) under irrigated condition.


However, the flag leaf width (0.114, -0.008), grain weight (0.315,-0.791)

and seed index (0.001, -0.020) were found to have opposite relationship between

correlation and their direct effect under irrigated condition, grain length (-0.383,

1.45) under TSD condition.

Beside these the character days to 50% flowering, (-0.001, -0.051) panicle

length (-0.018, -0.049), flag leaf area (-0.124, 0.012) and grain L: B ratio (0.372, -

1.731) showed vice-versa relationship between direct effect and correlation

coefficient for yield.

Thus in QTL, DTY 1.1 & 3.2 lines consider only those indirectly affecting

characters which enhances the value of correlation which ultimately enhances

yield.

4.4.3 Path analysis in QTL DTY 3.2 lines

Path analysis of different characters contributing towards yield revealed

that flag leaf length (0.251, 0.215), flag leaf width (0.241, 0.174), grain length

(0.133, 0.383) biological yield (0.735, 0.559,) and harvest index (0.712, 0.673)

showed positive direct effects while the characters Flag leaf area (-0.343, -0.305)

grain width (-0.153, -0.036) Grain L:B ratio (-0.201, -0.523) showed negative

direct effect in both conditions. The remaining characters vice versa effect on

either of the conditions viz. TSD and irrigated conditions (Table 4.11).

Among the positive direct effects high effect was show by biological yield

per plant (0.735, 0.559) followed to harvest Index (0.712, 0.673.) and grain length

(0.133, 0.383) the character biological yield (0.735, 0.559,) was found to be highly

significant in both the conditions with yield. Harvest Index also showed positive

correlation with grain yield hence during selection course the characters should be
given emphasis and the breeder should go for direct selection of these traits for

improving the yielding potential. The character grain length (0.133, 0.383) can

also be given due emphasis for direct as the association with grain yield is found

positive in both the condition.

Beside those the we can also go for direct selection of characters like

panicle length (0.026, -0.041), flag leaf length (0.251, 0.215), flag leaf area (-

0.343, -0.305), grain L: B ratio (-0.201, -0.523), grain weight (0.315,-0.791) and

seed index (0.001, -0.020) keeping in view of their association with grain yield.

4.4.4 Path analysis in without QTL lines

In without QTL lines correlation coefficients of various characters were

spliced out into direct and indirect affects both the condition (Table 4.12). Some

character exert positive direct effect like plant height (0.044, 0.003), flag leaf area

(0.068, 0.300), grain breadth (0.112, 0.208), grain L:B ratio (0.125, 0.236),

biological yield (0.820, 0.628) and harvest index (0.795, 0.562) while some exert

negative direct effect like days to 50% flowering ( -0.002, -0.067) flag leaf length

(-0.129, -0.268) flag leaf width (-0.056, -0.197) grain length (-0.071, -0.054)

and Seed index (-0.023, -0.051), while vice- versa effect were exhibited by

character panicle length ( 0.011, -0.027).

This indicates that direct selection of traits plant height (0.044, 0.003)

biological yield (0.820, 0.628), harvest index (0.795, 0.562) grain breadth (0.112,

0.208) days to 50% flowering (-0.002, -0.067), flag leaf length (-0.129, -0.268)

should be given more emphasis as the exert direct effect with yield under both

condition.
Beside these the character, Panicle length, Grain length, grain L:B ratio and

Seed index showed vice-versa relationship between direct effect and correlation

coefficient for yield. Hence, one should go for selection of such trait via desirable

indirect effects.

Overall the presence/absence of DT QTLs has changed the direct and

indirect effect of different characters to some extent, but the overall sscenario

remains same. Biological yield had significant direct effect and exhibited positive

correlation with grain yield under both the conditions, then in selection this

character should be given due importance.

The above findings for all the clusters for path analysis was in accordance

with finding of Chakraborty et al. (2001), Khedikar et al. (2004), Bisne et al.

(2006)and Manomani and Ranganath (2006) for character Seed Index for the

cluster QTL DTY 1.1, QTL DTY 1.1 and QTL DTY 3.2 under Irrigated condition

and QTL DTY 1.1 and 3.2 and Check lines under TSD condition; Mishra and

Verma (2002), Naik et al. (2005) and Panwar and Ali (2007) for Biological yield

under both condition for all the clusters except Check lines under TSD condition;

Khedikar et al. (2004), Manomani and Ranganathan (2006) for Days to 50%

Flowering for the cluster check lines under both condition and QTL DTY 3.2 for

Irrigated condition; Naik et al. (2005) for the character Grain length under both

condition for the QTL DTY 3.2 lines, irrigated condition for the QTL DTY 1.1 and

3.2 lines and Check lines, TSD condition for the QTL DTY line 1.1; vice versa for

different clusters for the grain breadth and Manomani and Ranganathan (2006) for

without QTL DTY lines under both condition for QTL DTY 3.2 under TSD

condition.
4.5 Effect of QTL for yield and yield contributing traits:

4.5.1 Performance of QTL over non QTL for days to 50% flowering:

Under terminal stage drought condition, all the lines i.e. with QTL 1.1,

with QTL 1.1 + 3.2 and with QTL 3.2 exhibited early days to 50% flowering as

compared to the lines without QTL. The mean ranged from 76 to 78 days. Apart

from this, there was at par performance in days to 50% flowering under irrigated

condition.

Table 4.13: Mean performance and percent change in days to 50% flowering
in lines with QTL and without QTL lines under Terminal stage
drought and irrigated condition.

With QTL1.1 With QTL 1.1 With QTL 3.2 Without QTL
(18) + 3.2 (16) (19) (19)
Terminal Stage Drought condition
Mean 78 76 76 80
% change -2.10 -4.12 -4.12 -
Irrigated condition
Mean 83 83 82 84
% change -1.40 -1.59 -2.1 -
Figure in parenthesis represent the number of lines.

4.5.2 Performance of QTL over non QTL for plant height:

The mean of plant height in TSD condition ranged from 120.9 cm (without

QTL) to 130.97 cm (With QTL 1.1+3.2). The results indicated high percent

change in plant height in lines with QTL 1.1+3.2 followed by lines with QTL 1.1

and with QTL 3.2. Furthermore, in irrigated condition all the lines showed high

means for plant height. The percent change for this trait ranged from 0.59 (with

QTL 3.2) to 0.67 (with QTL 1.1) so; QTL 1.1 is responsible to increase the plant

height (Table 4.14).


Table 4.14: Mean performance and percent change in Plant height in lines
with QTL and without QTL lines under Terminal stage drought
and irrigated condition.

With QTL1.1 With QTL 1.1 With QTL 3.2 Without QTL
(18) + 3.2 (16) (19) (19)
Terminal Stage Drought condition
Mean 129.32 130.97 123.47 120.9
% change 6.96 8.32 2.09 -
Irrigated condition
Mean 154.38 151.18 142.91 142.06
% change 8.67 6.42 0.598 -
Figure in parenthesis represent the number of lines.
4.5.3 Performance of QTL over non QTL for panicle length:

Under terminal stage drought condition, the mean of panicle length ranged

from 23.38 cm (with QTL 3.2) to 25.17 cm (with QTL 1.1). The results indicated

high percent change in panicle length in lines with QTL 1.1 followed by lines with

QTL 3.2 and lines with QTL 1.1+3.2. In irrigated condition, all the lines showed

high means for panicle length. The percent change for this trait ranged from 0.19

(with QTL 1.1+3.2) to 3.42 (with QTL 1.1) (Table 4.15).

Table 4.15: Mean performance and percent change in Panicle length in lines
with QTL and without QTL lines under Terminal stage drought
and irrigated condition.

With QTL1.1 With QTL 1.1 With QTL 3.2 Without QTL
(18) + 3.2 (16) (19) (19)
Terminal Stage Drought condition
Mean 25.17 23.62 23.38 23.68
% change 6.47 -0.08 -1.10 -
Irrigated condition
Mean 25.98 25.17 25.24 25.12
% change 3.42 0.19 0.47
Figure in parenthesis represent the number of lines.

4.5.4 Performance of QTL over non QTL for flag leaf length:
The mean of flag leaf length in terminal stage drought condition ranged

from 39.27 cm (without QTL) to 46.07 cm (with QTL 1.1+3.2). The results

exhibited high percent change in flag leaf length in lines with QTL 1.1+3.2

followed by lines with QTL 1.1 and with QTL 3.2. Furthermore, in irrigated

condition, all the lines exhibited lower mean of flag leaf length of with QTL and

without QTL lines. Then percent change for this trait ranged from 0.52 (with QTL

3.2) to 7.00 (with QTL 1.1+3.2) (Table 4.16).

Table 4.16: Mean performance and percent change in Flag leaf length in lines
with QTL and without QTL lines under Terminal stage drought
and irrigated condition.
With QTL 1.1 With QTL 1.1 With QTL 3.2 Without QTL
(18) + 3.2 (16) (19) (19)
Terminal Stage Drought condition
Mean 43.76 46.07 40.55 39.27
% change 11.43 17.31 3.25 -
Irrigated condition
Mean 37.84 38.64 36.11 35.92
% change 5.34 7.00 0.52 -
Figure in parenthesis represent the number of lines.

4.5.5 Performance of QTL over non QTL for flag leaf width:

The mean value for flag leaf width in TSD ranged from 1.19cm (with QTL

1.1) to 1.26 cm (with QTL 3.2). The results exhibited high percent change in flag

leaf width in lines with QTL 3.2 followed by lines with QTL 1.1. Furthermore, in

irrigated condition all the lines exhibited high means for flag leaf width. The

percent change for this trait ranged from (-) 4.31 (with QTL 1.1+3.2) to (-) 1.43

(with QTL 1.1) (Table 4.17).


Table 4.17: Mean performance and percent change in Flag leaf width in lines
with QTL and without QTL lines under Terminal stage drought
and irrigated condition.

With QTL 1.1 With QTL 1.1 + With QTL 3.2 Without QTL
(18) 3.2 (16) (19) (19)
Terminal Stage Drought condition
Mean 1.19 1.24 1.26 1.24
% change -4.03 0.00 1.61 -
Irrigated condition
Mean 1.37 1.3 1.39 1.39
% change -1.43 -4.31 0.00 -
Figure in parenthesis represent the number of lines.

4.5.6 Performance of QTL over non QTL for flag leaf area:

The mean of flag leaf area in TSD condition ranged from 34.07 cm2

(without QTL) to 39.31cm2 (with QTL 1.1+3.2). In irrigated condition, all the lines

showed lower mean for flag leaf area. The results exhibited high percent change

for this trait in lines with QTL 1.1+3.2 under both the condition i.e. terminal stage

drought and irrigated (Table 4.18).

Table 4.18: Mean performance and percent change in flag leaf area in lines
with QTL and without QTL lines under Terminal stage drought
and irrigated condition.

With QTL 1.1 With QTL 1.1 With QTL 3.2 Without QTL
(18) + 3.2 (16) (19) (19)
Terminal Stage Drought condition
Mean 35.89 35.87 35.21 34.92
% change 2.72 2.71 0.83
Irrigated condition
Mean 36.21 39.31 35.55 34.07
% change 6.28 15.38 4.34 -
Figure in parenthesis represent the number of lines.

4.5.7 Performance of QTL over non QTL for grain length:


The mean value of grain length in TSD condition ranged from 7.92 mm

(with QTL 1.1+3.2) to 8.32 mm (with QTL 3.2). The results indicated high percent

change in grain length in lines with QTL 3.2 followed by lines with QTL 1.1+3.2

and with QTL 1.1. Apart from this, in irrigated condition all the lines showed high

mean for grain length. The percent change for this trait ranged from (-) 1.83 (with

QTL 1.1) to (-) 2.71 (with QTL 1.1+3.2) (Table 4.19).

Table 4.19: Mean performance and percent change in Grain length in lines
with QTL and without QTL lines under Terminal stage drought
and irrigated condition.

With QTL 1.1 With QTL 1.1 With QTL 3.2 Without QTL
(18) + 3.2 (16) (19) (19)
Terminal Stage Drought condition
Mean 7.97 7.92 8.32 8.12
% change -1.78 -2.46 2.52 -
Irrigated condition
Mean 8.31 8.23 8.24 8.46
% change -1.83 -2.71 -2.66 -
Figure in parenthesis represent the number of lines.

4.5.8 Performance of QTL over non QTL for grain width:

The mean value of grain width in TSD condition ranged from 2.66 mm

(with QTL 1.1+3.2) to 2.75 mm (with QTL 3.2). The results indicated high percent

change in grain width in lines with QTL 3.2 followed by lines with QTL 1.1+3.2

and with QTL 1.1. Apart from this, in irrigated condition lines with QTL 1.1+3.2

showed high mean for grain width. The percent change for this trait ranged from (-

) 1.28 (with QTL 1.1) to 0.73 (with QTL 1.1+3.2) (Table 4.20).
Table 4.20: Mean performance and percent change in Grain width in lines
with QTL and without QTL lines under Terminal stage drought
and irrigated condition.

With QTL 1.1 With QTL 1.1 + With QTL 3.2 Without QTL
(18) 3.2 (16) (19) (19)
Terminal Stage Drought condition
Mean 2.73 2.66 2.76 2.75
% change -0.72 -3.21 0.36 -
Irrigated condition
Mean 2.69 2.75 2.72 2.73
% change -1.28 0.73 -0.36 -
Figure in parenthesis represent the number of lines.

4.5.9 Performance of QTL over non QTL for grain L: B ratio:

The average value of grain L: B ratio under TSD condition ranged from

2.94 (with QTL 1.1) to 3.04 (with QTL 3.2). The results indicated high percent

change in L: B ratio in lines with QTL 3.2 followed by lines with QTL 1.1+3.2 and

with QTL 1.1. Apart from this in irrigated condition all the lines exhibited high

means for grain L: B ratio. The percent change for this trait ranged from (-) 7.09

(with QTL 1.1+3.2) to (-) 4.01 (with QTL 1.1) (Table 4.21).

Table 4.21: Mean performance and percent change in grain L: B ratio in lines
with QTL and without QTL lines under Terminal stage drought
and irrigated condition.

With QTL 1.1 With QTL 1.1 + With QTL 3.2 Without QTL
(18) 3.2 (16) (19) (19)
Terminal Stage Drought condition
Mean 2.94 3.00 3.04 2.98
% change -1.34 0.67 2.01 -
Irrigated condition
Mean 3.11 3.01 3.04 3.24
% change -4.01 -7.09 -6.17
Figure in parenthesis represent the number of lines
4.5.10 Performance of QTL over non QTL for seed index:

The presence and absence of QTL did not make any difference under both

the condition for seed index (Table 4.22).

Table 4.22: Mean performance and percent change in Seed Index in lines with
QTL and without QTL lines under Terminal stage drought and
irrigated condition.

With QTL 1.1 With QTL 1.1 + With QTL 3.2 Without QTL
(18) 3.2 (16) (19) (19)
Terminal Stage Drought condition
Mean 1.95 2.04 2.16 2.11
% change -7.58 -1.89 2.37 --
Irrigated condition
Mean 2.14 2.18 2.20 2.17
% change -1.38 0.46 1.38 -
Figure in parenthesis represent the number of lines.

4.5.11 Performance of QTL over non QTL for harvest index:

The presence and absence of QTL did not make any difference under both

the condition for harvest index (Table 4.23).

Table 4.23: Mean performance and percent change in Harvest Index in lines
with QTL and without QTL lines under Terminal stage drought
and irrigated condition.

With QTL 1.1 With QTL 1.1 + With QTL 3.2 Without QTL
(18) 3.2 (16) (19) (19)
Terminal Stage Drought condition
Mean 28.44 26.59 28.87 27.07
% change 5.06 -1.77 6.64 -
Irrigated condition
Mean 21.28 21.64 22.59 23.00
% change -7.47 -5.91 -1.78 -
Figure in parenthesis represent the number of lines.

4.5.12 Performance of QTL over non QTL for biological yield:

The percent change in biological yield in the lines carrying QTLs indicates

the effect of QTL and biological yield is directly and significantly associated with

grain yield therefore, the QTL imparts advantage by increasing the biological yield

which ultimately also increases grain yield, but this advantage is prominent under

stress condition. While under irrigated situation the QTL leads to increase in plant

height responsible for increase in biological yield therefore, the lines carrying QTL

usually lodges (Table 4.24).

Table 4.24: Mean performance and percent change in Biological yield in lines
with QTL and without QTL lines under Terminal stage drought
and irrigated condition.

With QTL 1.1 With QTL 1.1 + With QTL 3.2 Without QTL
(18) 3.2 (16) (19) (19)
Terminal Stage Drought condition
Mean 439.81 430.21 423.69 338.87
% change 29.78 26.95 25.03 -
Irrigated condition
Mean 2090.12 1954.12 1925.74 1797.07
% change 16.30 8.74 7.16 -
Figure in parenthesis represent the number of lines.

4.5.13 Performance of QTL over non QTL for grain yield:

In TSD condition, the mean of grain yield varied from 103gm2 (without

QTL) to 122gm2 (with QTL 1.1 and QTL 3.2). Thus, the percent change in grain

yield also showed similar trend. As far as irrigated condition is concerned, the

grain yield showed at par performance.


This indicates clearly that presence of QTL imparts advantage to the line

only under stress condition and it has no advantage/ disadvantage under irrigated

condition. The presence of QTL in general increase plant height and therefore, the

lines with QTL under irrigated condition exhibit lodging which becomes

ultimately a disadvantage (Table 4.25).

Table 4.25: Mean performance and percent change in Grain yield in lines
with QTL and without QTL lines under terminal stage drought
and irrigated condition.

With QTL 1.1 With QTL 1.1 + With QTL 3.2 Without QTL
(18) 3.2 (16) (19) (19)
Terminal Stage Drought condition
Mean 122 116 122 103
% change 18 13 18 -
Irrigated condition
Mean 439 421 448 426
% change 3.1 -1.2 5.2 -
Figure in parenthesis represent the number of lines.

Quantitative trait loci (QTL) with large effects on yield under drought

stress have been detected in new populations derived from parents differing greatly

in yield under drought, using selective genotyping. In a Vandana/Way Rarem

population, a QTL on chromosome 12 (DTY12.1) explained 51% of the genetic

variance for yield under stress (Bernier et al. 2007). Another QTL on chromosome

4 (DTY4.1) affects yield under reproductive-stage stress in 1R55419-04/Way

Rarem population, where it explained over 35% of genetic variance (IRRI,

unpublished). The effect of QTL DTY12.1 has been validated in diverse

environments. However, these major QTLs showed low and variable effect in the

background of improved mega varieties, the targeted cultivars for improving

drought tolerance. Recently, two major QTLs, one each in the background of mega
varieties „Swarna‟ and „IR 64‟ have been identified. In Apo/2*Swarna F4.5

population, a major QTL (IJTY3.1) has been identified on chromosome 3 near

RM520 in the marker interval RM520-RM416. This QTL explained 31% of the

genetic variance of the trait (P<0.0001), and showed an average additive effect of

180 kg ha which accounted for 28% of the trial mean under stress (Venuprasad et

al. 2008). Similarly, in 1R77298-14-1-2/JR 64 population, another major QTL

(DTY2.1) has been identified on chromosome 2 for grain yield under reproductive

stage lowland drought stress near marker RM236. This QTL explained 77% of the

genetic variance of the trait (p<O.000I) and showed an average additive effect of

100 kg ha‟ (IRRI, unpublished). Recent work at IRRI has identified a robust QTL

for yield under drought on rice chromosome 1 (DTY1.1) which is effective across

the genetic backgrounds of 1R64, Swarna and Sambha Mahsuri and this has been

choosen for transfer into mega varieties.

The study also indicates that DT QTLs impart advantage to the line in

terms of increased grain yield under water limited environment.


CHAPTER-V

SUMMARY, CONCLUSION AND SUGGESTIONS FOR

FUTURE RESEARCH WORK

Among the abiotic stresses drought stress is a major constrain to stability in

production and yield under rainfed rice ecosystem. It acts as a serious limiting

factor in agriculture production by preventing a crop from reaching the genetically

determined theoretical maximum yield. The development of cultivars with

improved drought tolerance is thus an important element in increasing productivity

and alleviating poverty in communities depended on rainfed ecosystem. The

complex nature of drought tolerance, genotype x environment interaction, lack of

understanding of inheritance of drought tolerance, poor understanding of

physiological basis of yield under water limited condition and difficulty of

effective drought tolerance screening complicate the development of drought

tolerance varieties. Therefore, the present study was undertaken to carry out the

genetic analysis for understanding the genetics basis of drought tolerance and

identification of QTLs for yield and yield contributing traits under water stress

condition.

Seventy seven lines of drought tolerant QTL population were evaluated

under irrigated and TSD condition during wet season 2010. The experimental

material for the present investigation comprised 77 lines derived from Nagina

22/Swarna cross including checks (MTU-1010, IR-36, N-22, Swarna & IR-64).

The 77 lines were evaluated under irrigated and terminal stage drought condition
for genetic analysis and generating phenotypic data to know the effect of different

with and without QTLs lines on yield and yield contributing traits.

Conclusions

1. Presence of QTL‟s has increased the plant height.

2. Presence of QTL has resulted increased yield under stress condition and this is a

major advantage.

3. The increase in the plant height because of DT QTL‟s under irrigated condition

results in lodging of these lines which become a major disadvantage.

Suggestions for future work:

 DT QTL and plant height, this linkage needs to be broken down.


“EFFECT OF DROUGHT TOLERANT QTLs ON YIELD AND YIELD
CONTRIBUTING TRAITS UNDER MANAGED WATER STRESS AS
WELL AS NON STRESS CONDITION”

By
Manoj Kumar Narang
Abstract

The complex nature of drought tolerance, genotype x environment


interaction, lack of understanding of inheritance of drought tolerance, poor
understanding of physiological basis of yield under water limited condition and
difficulty of effective drought tolerance screening complicate the development of
drought tolerance varieties. Therefore, the present study was undertaken to carry
out the genetic analysis for understanding the genetics basis of drought tolerance
and identification of QTLs for yield and yield contributing traits under water stress
condition.
The analysis of variance indicated substantial variation among the 77 lines
for grain yield and yield contributing traits. Significant variation was noticed
among the 77 lines for grain yield and yield contributing traits under irrigated as
well as TSD situation. The mean grain yield of these lines was higher in irrigated
as compared to TSD condition, respectively, indicating very high level of stress.
The presence of QTL‟s has increased the plant height and also it has resulted
increased yield under stress condition which is a major advantage.. The increase in
the plant height because of DT QTL‟s under irrigated condition results in lodging
of these lines which become a major disadvantage. The estimates of correlation
showed highly positive and significant association between grain yield, harvest
index and biological yield under both irrigated as well as terminal stage drought
condition. The estimates of direct effect showed that biological yield had
significant direct effect and exhibited positive correlation with grain yield under
both the conditions.
The major QTLs showed low and variable effect in the background of
improved mega varieties, the targeted cultivars for improving drought tolerance.
Recently, two major QTLs, one each in the background of mega varieties „Swarna‟
and „IR 64‟ have been identified.

(Dr. S.B. Verulkar)


Department of Genetics and Plant Breeding Chairman, Advisory Committee
College of Agriculture, IGKV, Raipur
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Table 4.1: Analysis of variance for different characters in rice under terminal
stage drought and irrigated condition during Kharif 2010

Mean sum of squares due to genotypes


Characters

Terminal stage drought Irrigated

Days to 50 per cent flowering 34.42** 43.33**

Plant height (cm) 526.65** 1032.85**

Panicle length (cm) 4.80 9.07

Flag leaf length (cm) 90.88** 45.83

Flag leaf Width (cm) 0.05 0.04

FLA - Flag leaf area (cm2) 84.94 59.72

Grain Length (mm) 3.67** 3.11**

Grain Width (mm) 0.27 0.25

Grain L:B ratio 0.13 0.24

100 seed weight (g) 0.14** 0.11**

Harvest index (%) 72.07** 78.07**

Biological yield (g/m2) 11575.65** 582672.84**

Grain yield (g/m2) 2047.62 69983.36**

* Significant at 0.05 probability level and ** = Significant at 0.01 probability


level
Table 4.2:MEAN OF GRAIN YIELD WITH DSI IN TSD AND IRRIGATED
CONDITION

TSD IRRIGATED TSD


E.NO MEAN MEAN YIELD DSI
DESIGNATION QTLs INFORMATION YIELD
1 IR 86931-B-6 with QTL DTY3.2 110.0 588.9 1.11
2 IR 86931-B-9 with QTL DTY3.2 133.3 588.9 1.06
3 IR 86931-B-26 with QTL DTY1.1 & 3.2 136.7 644.4 1.08
4 IR 86931-B-32 with QTL DTY3.2 101.0 477.8 1.08
5 IR 86931-B-36 without any QTL 123.3 466.7 1.00
6 IR 86931-B-40 without any QTL 116.7 255.6 0.74
7 IR 86931-B-50 without any QTL 136.7 400.0 0.90
8 IR 86931-B-55 with QTL DTY1.1 & 3.2 116.7 388.9 0.96
9 IR 86931-B-56 with QTL DTY1.1 & 3.2 123.3 588.9 1.08
10 IR 86931-B-60 with QTL DTY3.2 150.0 500.0 0.96
11 IR 86931-B-66 with QTL DTY1.1 156.7 377.8 0.80
12 IR 86931-B-67 with QTL DTY3.2 93.3 455.6 1.09
13 IR 86931-B-78 without any QTL 116.7 488.9 1.04
14 IR 86931-B-86 with QTL DTY1.1 136.7 388.9 0.89
15 IR 86931-B-103 without any QTL 103.3 344.4 0.96
16 IR 86931-B-104 without any QTL 90.0 155.6 0.58
17 IR 86931-B-120 without any QTL 113.3 411.1 0.99
18 IR 86931-B-131 with QTL DTY1.1 146.7 377.8 0.84
19 IR 86931-B-132 with QTL DTY3.2 163.3 766.7 1.07
20 IR 86931-B-147 with QTL DTY1.1 110.0 444.4 1.03
21 IR 86931-B-150 with QTL DTY1.1 110.0 522.2 1.08
22 IR 86931-B-152 without any QTL 170.0 422.2 0.82
23 IR 86931-B-158 without any QTL 50.0 200.0 1.02
24 IR 86931-B-142 without any QTL 123.3 344.4 0.88
25 IR 86931-B-166 with QTL DTY1.1 & 3.2 116.7 433.3 1.00
26 IR 86931-B-170 with QTL DTY3.2 150.0 388.9 0.84
27 IR 86931-B-173 without any QTL 113.3 388.9 0.97
28 IR 86931-B-187 with QTL DTY1.1 65.0 300.0 1.07
29 IR 86931-B-192 with QTL DTY1.1 156.7 855.6 1.12
30 IR 86931-B-194 without any QTL 93.3 300.0 0.94
31 IR 86931-B-224 with QTL DTY1.1 150.0 255.6 0.56
32 IR 86931-B-233 with QTL DTY1.1 120.0 500.0 1.04
33 IR 86931-B-256 with QTL DTY1.1 & 3.2 80.0 377.8 1.08
34 IR 86931-B-263 without any QTL 76.7 233.3 0.92
35 IR 86931-B-279 with QTL DTY1.1 & 3.2 113.3 1022.2 1.21
36 IR 86931-B-294 with QTL DTY1.1 & 3.2 100.0 522.2 1.10
37 IR 86931-B-301 with QTL DTY1.1 166.7 777.8 1.07
38 IR 86931-B-311 without any QTL 106.7 477.8 1.06
39 IR 86931-B-323 with QTL DTY1.1 130.0 688.9 1.11
40 IR 86931-B-326 without any QTL 120.0 477.8 1.02
41 IR 86931-B-310 with QTL DTY1.1 113.3 400.0 0.98
42 IR 86931-B-346 with QTL DTY1.1 156.7 222.2 0.40
43 IR 86931-B-347 with QTL DTY1.1 & 3.2 93.3 88.9 -0.07
44 IR 86931-B-365 with QTL DTY3.2 110.0 611.1 1.12
45 IR 86931-B-366 with QTL DTY3.2 153.3 733.3 1.08
46 IR 86931-B-373 with QTL DTY1.1 & 3.2 146.7 333.3 0.76
47 IR 86931-B-390 with QTL DTY3.2 140.0 422.2 0.91
48 IR 86931-B-400 with QTL DTY1.1 123.3 333.3 0.86
49 IR 86931-B-414 with QTL DTY1.1 & 3.2 106.7 444.4 1.04
50 IR 86931-B-431 without any QTL 36.7 233.3 1.15
51 IR 86931-B-454 with QTL DTY3.2 193.3 344.4 0.60
52 IR 86931-B-456 with QTL DTY1.1 & 3.2 128.3 122.2 -0.07
53 IR 86931-B-464 with QTL DTY3.2 80.0 255.6 0.94
54 IR 86931-B-467 with QTL DTY3.2 93.3 455.6 1.09
55 IR 86931-B-486 with QTL DTY3.2 126.7 555.6 1.05
56 IR 86931-B-487 without any QTL 90.0 766.7 1.20
57 IR 86931-B-498 with QTL DTY1.1 91.7 222.2 0.80
58 IR 86931-B-502 with QTL DTY1.1 & 3.2 90.0 266.7 0.90
59 IR 86931-B-516 with QTL DTY1.1 & 3.2 70.0 700.0 1.23
60 IR 86931-B-519 with QTL DTY1.1 & 3.2 96.7 755.6 1.19
61 IR 86931-B-17 with QTL DTY1.1 93.3 422.2 1.06
62 IR 86931-B-521 with QTL DTY1.1 96.7 544.4 1.12
63 IR 86931-B-523 with QTL DTY3.2 120.0 355.6 0.90
64 IR 86931-B-524 with QTL DTY1.1 & 3.2 113.3 188.9 0.55
65 IR 86931-B-547 with QTL DTY3.2 73.3 300.0 1.03
66 IR 86931-B-549 with QTL DTY3.2 68.3 288.9 1.04
67 IR 86931-B-552 with QTL DTY1.1 & 3.2 106.7 300.0 0.88
68 IR 86931-B-559 without any QTL 96.7 233.3 0.80
69 IR 86931-B-561 with QTL DTY3.2 130.0 800.0 1.14
70 IR 86931-B-564 with QTL DTY1.1 96.7 233.3 0.80
71 IR 86931-B-574 with QTL DTY3.2 130.0 255.6 0.67
72 IR 86931-B-578 without any QTL 173.3 455.6 0.85
73 N22 Check 163.3 288.9 0.59
74 Swarna Check 10.0 211.1 1.30
75 IR 64 Check 140.0 544.4 1.01
76 IR 36 Check 118.3 322.2 0.86
77 MTU 1010 Check 136.7 655.6 1.08
Drought Intensity Index (DII) = (1-Xs/Xi) (Ramírez-Vallejo and Kelly,1998)
DSI = (1-Ys/Yi)/DII (Fischer and Maurer, 1978)
Table 4.3: Genetics parameters of variation for yield and its components under TSD
and Irrigated condition (Kharif 2010)

Character Condition Mean Range GCV PCV h2 GA GA %


max min (%) mean
DF TSD 77.79 91.50 67.00 5.25 5.42 0.93 8.15 10.51
Irrigated 82.84 97.00 71.00 5.61 5.63 0.99 9.54 11.51
PH TSD 123.90 150.80 75.65 12.49 13.68 0.83 29.09 23.47
Irrigated 144.83 176.00 79.65 14.38 16.90 0.72 36.51 25.20
PL TSD 23.81 28.85 17.20 4.27 7.53 0.32 1.26 4.97
Irrigated 25.31 29.20 20.50 5.67 11.30 0.25 1.40 5.87
FLL TSD 41.61 55.00 28.05 13.11 18.78 0.48 7.85 18.86
Irrigated 36.59 49.00 27.05 6.25 17.41 0.12 1.69 4.61
FLW TSD 1.23 1.65 0.85 6.90 17.22 0.16 0.07 5.69
Irrigated 1.37 1.70 1.10 2.30 15.05 0.02 0.01 0.72
FLA TSD 35.70 51.45 23.35 11.83 22.94 0.26 4.49 12.57
Irrigated 35.12 45.25 25.70 5.05 21.42 0.05 0.86 2.44
GL TSD 8.14 9.93 6.68 7.06 9.42 0.56 1.77 10.64
Irrigated 8.34 10.03 7.43 5.95 8.74 0.46 1.39 8.33
GW TSD 2.73 3.10 2.05 3.30 8.98 0.13 0.14 2.56
Irrigated 2.69 3.15 1.95 3.34 8.68 0.14 0.14 2.61
LB TSD 3.00 3.77 2.51 5.58 10.94 0.26 0.18 6.00
Irrigated 3.12 4.01 2.12 7.11 14.16 0.25 0.23 7.37
SI TSD 2.07 2.54 1.49 10.00 15.02 0.44 0.28 13.52
Irrigated 2.17 2.60 1.62 8.80 13.12 0.44 0.26 11.98
HI TSD 27.82 39.20 3.00 19.11 23.79 0.64 8.80 31.63
Irrigated 22.12 43.32 10.98 24.61 31.44 0.61 8.78 39.69
BY TSD 416.71 566.65 133.35 16.83 19.58 0.73 124.10 29.78
Irrigated 1928.85 3322.00 788.90 24.61 30.99 0.63 776.44 40.25
GY TSD 115.53 173.35 10.00 25.77 29.49 0.76 53.59 46.39
Irrigated 432.02 1022.20 122.20 39.31 46.94 0.70 293.02 67.82

DF =Days to 50 per cent flowering; PH= Plant height (cm); PL =Panicle length (cm); FLL =Flag leaf
length (cm); FLW =Flag leaf Width (cm); FLA=Flag leaf area (cm2); GL =Grain Length (mm); GW =Grain
Width (mm); LB =Grain L:B ratio; SI =100 seed weight (g); HI =Harvest index (%);BY =Biological yield
(g/m2); GY =Grain yield (g/m2); TSD = Terminal Stage Drought
Table 4.5: Correlation coefficient of grain yield and its component in with QTL DTY 1.1 & 3.2 line

PH PL FLL FLW FLA GL GW LB SI HI BY GY


T 0.049 -0.367 0.431 -0.113 0.247 0.152 0.134 -0.001 0.144 -0.020 0.193 0.063
DF -0.093 0.213 -0.186 0.287 0.128 0.101 0.149 0.020 0.234 0.323 0.192 0.285
I
T -0.079 0.569* -0.243 0.300 -0.613* 0.020 -0.511* -0.042 -0.145 0.243 -0.026
PH
I 0.005 0.409 -0.064 0.204 -0.337 0.120 -0.356 -0.240 -0.566* 0.286 -0.057
T 0.075 0.184 0.123 -0.015 -0.007 0.028 -0.079 0.071 0.183 0.137
PL
I 0.181 0.355 0.409 0.177 0.320 0.002 0.343 0.367 0.186 0.340
T -0.363 0.566* -0.477* 0.012 -0.401 -0.122 -0.068 0.368 0.106
FLL
I 0.001 0.595* -0.253 0.130 -0.278 0.343 0.367 0.186 0.340
T 0.545* 0.218 -0.154 0.153 0.128 -0.065 0.304 0.076
FLW
I 0.792** 0.111 0.073 0.079 0.343 0.367 0.186 0.340
T -0.254 -0.077 -0.150 0.153 0.128 -0.065 0.076
FLA
I -0.062 0.141 -0.107 0.201 -0.083 0.045 -0.005
T 0.152 0.192 -0.057 0.151 -0.187 0.007
GL
I -0.078 0.883** 0.163 0.302 0.094 0.224
T -0.597* 0.237 0.099 0.201 0.164
GW
I -0.534* 0.253 0.034 0.167 0.173
T 0.342 0.066 -0.290 -0.097
LB
I 0.024 0.245 -0.011 0.097
T 0.148 -0.120 0.017
SI
I 0.483* 0.164 0.381
HI T 0.242 0.871**
I -0.054 0.483*
T 0.676**
BY
I 0.826**

DF - Days to 50 per cent flowering PH - Plant height (cm) PL - Panicle length (cm) FLL – Flag leaf length (cm),
FLW - Flag leaf Width (cm) FLA - Flag leaf area (cm2) GL –Grain Length (mm) GW – Grain Width (mm)
LB – Grain L: B ratio SI -100 seed weight (g) HI-Harvest index (%) BY -Biological yield (g/m2)
GYP - Grain yield (g/m2) T – TSD condition I – Irrigated condition

* Significant at 5% level of significance and **Significant at 1% level of significance


Table 4.10: Path coefficient analysis showing direct and indirect effects of different yield contributing character on yield in with
QTL DTY 1.1 & 3.2 lines
DF PH PL FLL FLW FLA GL GW LB SI HI BY

T -0.001 -0.003 0.007 0.038 -0.013 -0.031 -0.058 0.042 0.000 0.000 -0.015 0.098
DF
I -0.051 0.004 -0.011 -0.017 -0.002 0.002 0.147 -0.118 -0.035 -0.005 0.195 0.171
T 0.000 -0.056 0.002 0.050 -0.028 -0.037 0.234 0.006 -0.190 0.000 -0.109 0.123
PH
I 0.005 -0.044 0.000 0.037 0.001 0.003 -0.489 -0.095 0.615 0.005 -0.342 0.254
T 0.000 0.005 -0.018 -0.007 0.021 -0.015 0.006 -0.002 0.010 0.000 0.054 0.093
PL
I -0.011 0.000 -0.049 0.016 -0.003 0.005 0.257 -0.253 -0.004 -0.007 0.222 0.165
T 0.000 -0.032 0.001 0.089 -0.042 -0.070 0.182 0.004 -0.149 0.000 -0.051 0.186
FLL
I 0.009 -0.018 -0.009 0.091 0.000 0.007 -0.367 -0.103 0.482 0.001 -0.245 0.028
T 0.000 0.014 -0.003 -0.032 0.114 -0.068 -0.083 -0.049 0.113 0.000 0.097 -0.033
FLW
I -0.015 0.003 -0.018 0.000 -0.008 0.010 0.160 -0.058 -0.137 -0.007 0.116 0.021
T 0.000 -0.017 -0.002 0.050 0.062 -0.124 0.097 -0.024 -0.056 0.000 0.023 0.149
FLA
I -0.007 -0.009 -0.020 0.054 -0.006 0.012 -0.090 -0.112 0.185 -0.004 -0.050 0.040
T 0.000 0.034 0.000 -0.042 0.025 0.032 -0.383 0.084 0.226 0.001 0.114 -0.095
GL
I -0.005 0.015 -0.009 -0.023 -0.001 0.001 1.451 0.062 -1.529 -0.003 0.183 0.084
T 0.000 -0.001 0.000 0.001 -0.018 0.010 -0.102 0.315 -0.222 0.000 0.075 0.102
GW
I -0.008 -0.005 -0.016 0.012 -0.001 0.002 -0.113 -0.791 0.925 -0.005 0.021 0.148
T 0.000 0.029 -0.001 -0.035 0.035 0.019 -0.233 -0.188 0.372 0.000 0.050 -0.147
LB
I -0.001 0.016 0.000 -0.025 -0.001 -0.001 1.281 0.423 -1.731 -0.001 0.148 -0.010
T 0.000 0.002 0.002 -0.011 0.017 0.001 -0.245 0.075 0.126 0.001 0.112 -0.061
SI
I -0.012 0.011 -0.017 -0.005 -0.003 0.003 0.236 -0.200 -0.042 -0.020 0.292 0.146
T 0.000 0.008 -0.001 -0.006 0.015 -0.004 -0.058 0.031 0.025 0.000 0.753 0.122
HI
I -0.016 0.025 -0.018 -0.037 -0.002 -0.001 0.439 -0.027 -0.423 -0.010 0.604 -0.048
T 0.000 -0.014 -0.003 0.033 -0.008 -0.037 0.071 0.063 -0.108 0.000 0.182 0.506
BY
I -0.010 -0.012 -0.009 0.003 0.000 0.001 0.137 -0.132 0.019 -0.003 -0.033 0.887
TSD RESIDUAL EFFECT =0.06 IRRIGATED RESIDUAL EFFECT =0.02

DF - Days to 50 per cent flowering PH - Plant height (cm) PL - Panicle length (cm) FLL – Flag leaf length (cm),
FLW - Flag leaf Width (cm) FLA - Flag leaf area (cm2) GL –Grain Length (mm) GW – Grain Width (mm)
LB – Grain L: B ratio SI -100 seed weight (g) HI-Harvest index (%) BY -Biological yield (g/m2)
T – TSD condition I – Irrigated condition
Table: 4.4: Correlation coefficient of grain yield and its component in with QTL DTY 1.1 line

PH PL FLL FLW FLA GL GW LB SI HI BY GY


T -0.016 0.052 -0.100 0.183 0.086 0.278 0.152 0.096 -0.203 0.070 -0.035 0.016
DF
I -0.016 0.052 -0.100 0.183 0.086 -0.060 0.188 -0.203 0.010 -0.052 -0.117 -0.191
T 0.040 0.424 -0.169 0.249 -0.100 0.047 -0.163 -0.317 -0.490* 0.181 -0.441
PH
I 0.073 0.075 0.028 0.095 -0.434 -0.396 0.152 -0.088 -0.553* 0.349 -0.146

T 0.008 0.093 0.094 0.034 -0.063 0.096 -0.317 -0.490* 0.181 -0.441
PL
I 0.299 0.245 0.448 0.197 -0.183 0.241 0.236 -0.062 0.312 0.165

T -0.308 0.525* -0.127 0.138 -0.259 -0.129 -0.111 -0.046 -0.153


FLL
I -0.263 0.626** -0.101 -0.150 0.040 -0.196 -0.099 0.490* 0.311

T 0.629** 0.110 -0.030 0.134 0.169 0.212 0.035 0.237


FLW
I 0.577* -0.046 0.189 -0.187 -0.047 0.059 -0.094 -0.067

T -0.022 0.119 -0.142 0.169 0.212 0.035 0.237


FLA I -0.096 0.0567 -0.131 -0.213 -0.049 0.364 0.220

T 0.454 0.399 0.211 0.096 -0.099 -0.042


GL
I 0.007 0.628** 0.668 0.292 0.049 0.257

T -0.634** 0.211 0.096 -0.099 -0.042


GW
I -0.765** 0.072 -0.112 0.154 0.005

T -0.158 -0.023 0.045 0.176


LB
I 0.072 -0.112 0.154 0.005
T 0.218 0.247 0.457
SI
I 0.174 0.124 0.216

T -0.608 0.697**
HI
I -0.170 0.624**

T 0.618**
BY
I 0.634**
DF - Days to 50 per cent flowering PH - Plant height (cm) PL - Panicle length (cm) FLL – Flag leaf length (cm),
FLW - Flag leaf Width (cm) FLA - Flag leaf area (cm2) GL –Grain Length (mm) GW – Grain Width (mm)
LB – Grain L: B ratio SI -100 seed weight (g) HI-Harvest index (%) BY -Biological yield (g/m2)
GY - Grain yield (g/m2) T – TSD condition I – Irrigated condition

* - Significant at 5% level of significance and ** - Significant at 1% level of significance


Table 4.9: Path coefficient analysis showing direct and indirect effects of different yield contributing character on yield in with
QTL DTY 1.1 lines
DF PH PL FLL FLW FLA GL GW LB SI HI BY

-0.021 0.000 0.001 -0.021 0.042 -0.023 0.048 -0.041 -0.038 0.008 0.087 -0.029
T
DF
I -0.032 0.001 0.009 -0.066 -0.018 0.081 0.020 0.047 -0.072 0.000 -0.042 -0.082
0.000 -0.011 0.001 0.089 -0.039 -0.068 -0.017 -0.012 0.064 0.012 -0.614 0.152
T
PH
I 0.001 -0.026 0.002 -0.030 -0.012 0.046 0.146 0.038 -0.141 -0.002 -0.448 0.244
-0.001 0.000 0.035 0.001 0.021 -0.025 0.005 0.017 0.038 -0.004 -0.005 0.063
T
PL
I -0.008 -0.001 0.033 -0.122 -0.106 0.221 -0.066 -0.045 0.085 0.005 -0.050 0.218
0.002 -0.004 0.000 0.211 -0.071 -0.144 -0.022 -0.037 0.102 0.005 -0.139 -0.038
T
FLL
I -0.005 -0.002 0.010 -0.411 0.113 0.308 0.034 -0.037 0.014 -0.004 -0.080 0.343
-0.004 0.001 0.003 -0.065 0.230 -0.172 0.019 0.008 -0.052 -0.006 0.266 0.029
T
FLW
I -0.001 0.000 0.008 0.108 -0.433 0.284 0.015 0.047 -0.066 -0.001 0.047 -0.065
-0.001 -0.002 0.003 0.111 0.145 -0.274 -0.003 -0.032 0.056 -0.003 0.093 0.003
T
FLA
I -0.005 -0.002 0.015 -0.257 -0.249 0.493 0.032 0.014 -0.046 -0.004 -0.040 0.255
-0.006 0.001 0.001 -0.027 0.025 0.006 0.174 -0.123 -0.157 -0.008 0.120 -0.083
T
GL
I 0.001 0.011 0.006 0.041 0.019 0.047 -0.338 0.001 0.223 0.015 0.237 0.034
-0.003 0.000 -0.002 0.029 -0.006 0.032 0.079 -0.273 0.250 -0.013 0.143 -0.041
T
GW
I -0.066 -0.004 -0.006 0.061 -0.082 0.027 -0.002 0.251 0.272 0.001 -0.091 0.107
-0.002 0.001 0.003 -0.054 0.030 0.038 0.069 0.173 -0.395 0.006 -0.028 -0.038
T
LB
I 0.006 0.010 0.008 -0.016 0.081 -0.064 -0.212 -0.192 0.356 0.008 0.227 -0.074
0.004 0.003 0.004 -0.027 0.039 -0.025 0.036 -0.091 0.062 -0.040 0.273 0.209
T
SI
I 0.000 0.002 0.008 0.080 0.020 -0.104 -0.225 0.018 0.136 0.022 0.141 0.086
-0.001 0.005 0.000 -0.023 0.049 -0.020 0.016 -0.031 0.009 -0.008 1.254 -0.513
T
HI
I 0.001 0.014 -0.002 0.040 -0.025 -0.024 -0.098 -0.028 0.099 0.003 0.811 -0.118
0.000 -0.002 0.002 -0.009 0.008 -0.001 -0.017 0.013 0.017 -0.010 -0.762 0.844
T
BY
I 0.003 -0.009 0.010 -0.201 0.040 0.179 -0.016 0.038 -0.037 0.002 -0.137 0.701
TSD RESIDUAL EFFECT = 0.02 IRRIGATED RESIDUAL EFFECT = 0.02

DF - Days to 50 per cent flowering PH - Plant height (cm) PL - Panicle length (cm) FLL – Flag leaf length (cm),
FLW - Flag leaf Width (cm) FLA - Flag leaf area (cm2) GL –Grain Length (mm) GW – Grain Width (mm)
LB – Grain L: B ratio SI -100 seed weight (g) HI-Harvest index (%) BY -Biological yield (g/m2)
T – TSD condition I – Irrigated condition
Table 4.6: Correlation coefficient of grain yield and its component in with QTL DTY 3.2 lines

PH PL FLL FLW FLA GL GW LB SI HI BY GY


T -0.071 -0.304 -0.048 0.283 0.135 -0.155 -0.153 0.010 -0.090 -0.333 0.004 -0.273
DF
I -0.071 -0.054 -0.219 -0.002 -0.207 -0.151 0.071 -0.143 -0.330 -0.004 0.043 0.064
T 0.425 0.349 -0.148 0.225 0.039 -0.200 0.222 -0.011 -0.072 0.489* 0.292
PH
I 0.218 0.289 -0.066 0.232 0.091 0.418 -0.257 0.379 -0.457 0.308 -0.144

T 0.185 -0.300 -0.045 0.061 -0.130 0.172 0.074 -0.093 0.206 0.109
PL
I 0.394 -0.019 0.313 -0.157 -0.014 -0.102 0.007 0.101 0.064 0.046

FLL T -0.155 0.716** -0.345 -0.059 -0.210 -0.308 -0.332 0.340 0.006

I -0.055 0.801** 0.067 0.037 0.009 0.066 -0.213 0.150 -0.119

FLW T 0.561* -0.040 -0.035 -0.006 0.027 -0.288* -0.054 -0.256

I 0.537* 0.047 -0.368 0.299 -0.126 0.199 0.230 0.257

FLA T -0.317 -0.107 -0.152 -0.249 -0.459 0.278 -0.143

0.075 -0.141 0.143 -0.074 0.275 0.048 0.008

GL T 0.307 0.501* 0.647** 0.263 -0.201 0.022

I 0.058 0.655** 0.305 0.088 0.110 0.146

GW T -0.665 0.562* 0.240 -0.059 0.146

I -0.714 0.424 -0.321 0.107 -0.122

LB T -0.020 0.004 -0.109 -0.108

I -0.119 0.314 -0.012 0.195


SI T 0.282 0.180 0.056

I -0.055 0.120 0.062

HI T -0.055 0.676**

I 0.221 0.801**

BY T 0.691**

I 0.672**
DF - Days to 50 per cent flowering PH - Plant height (cm) PL - Panicle length (cm) FLL – Flag leaf length (cm),
FLW - Flag leaf Width (cm) FLA - Flag leaf area (cm2) GL –Grain Length (mm) GW – Grain Width (mm)
LB – Grain L: B ratio SI -100 seed weight (g) HI-Harvest index (%) BY -Biological yield (g/m2)
GY - Grain yield (g/m2) T – TSD condition I – Irrigated condition

* Significant at 5% level of significance and **Significant at 1% level of significance


Table 4.11: Path coefficient analysis showing direct and indirect effects of different yield contributing character on yield in with
QTL DTY 3.2 lines

DF PH PL FLL FLW FLA GL GW LB SI HI BW

T -0.046 0.000 -0.008 -0.012 0.068 -0.046 -0.020 0.023 -0.002 0.003 -0.236 0.003
DF
I 0.039 0.000 0.002 -0.047 0.000 0.063 -0.057 -0.025 0.074 -0.011 -0.002 0.023
T 0.003 0.000 0.011 0.087 -0.035 -0.077 0.005 0.030 -0.044 0.000 -0.051 0.359
PH
I -0.002 -0.008 -0.009 0.062 -0.011 -0.070 0.034 -0.150 0.134 0.013 -0.307 0.171
T 0.014 0.000 0.026 0.046 -0.072 0.015 0.008 0.019 -0.034 -0.002 -0.066 0.151
PL
I -0.002 -0.001 -0.041 0.084 -0.003 -0.095 -0.060 0.004 0.053 0.001 0.068 0.035
T 0.002 0.000 0.004 0.251 -0.037 -0.246 -0.045 0.009 0.042 0.010 -0.236 0.249
FLL
I -0.008 -0.002 -0.016 0.215 -0.009 -0.244 0.025 -0.013 -0.004 0.002 -0.143 0.084
T -0.013 0.000 -0.007 -0.039 0.241 -0.193 -0.005 0.005 0.001 -0.001 -0.205 -0.039
FLW
I 0.000 0.001 0.001 -0.011 0.174 -0.164 0.018 0.132 -0.156 -0.004 0.133 0.128
T -0.006 0.000 -0.001 0.179 0.135 -0.343 -0.042 0.016 0.030 0.008 -0.326 0.204
FLA
I -0.008 -0.002 -0.013 0.172 0.093 -0.305 0.028 0.051 -0.074 0.001 -0.049 0.153
T 0.007 0.000 0.001 -0.086 -0.009 0.109 0.133 -0.047 -0.100 -0.022 0.187 -0.147
GL
I -0.006 -0.001 0.006 0.014 0.008 -0.023 0.383 -0.020 -0.342 0.010 0.059 0.061
T 0.007 0.000 -0.003 -0.014 -0.008 0.036 0.040 -0.153 0.133 -0.019 0.170 -0.043
GW
I 0.002 -0.003 0.001 0.008 -0.064 0.043 0.022 -0.036 0.373 0.015 -0.216 0.059
T 0.000 0.000 0.004 -0.052 -0.001 0.052 0.066 0.102 -0.201 0.000 0.002 -0.080
LB
I -0.005 0.002 0.004 0.002 0.052 -0.043 0.251 0.257 -0.523 -0.004 0.211 -0.006
T 0.004 0.000 0.002 -0.077 0.006 0.085 0.086 -0.086 0.004 -0.035 0.200 -0.132
SI
I -0.013 -0.003 -0.001 0.014 -0.022 -0.002 0.117 -0.153 0.062 0.035 -0.036 0.066
T 0.015 0.000 -0.002 -0.083 -0.069 0.157 0.035 -0.036 0.000 -0.010 0.712 -0.040
HI
I -0.002 0.003 -0.004 -0.045 0.034 0.022 0.033 0.115 -0.164 -0.001 0.673 0.123
T 0.000 0.000 0.005 0.085 -0.013 -0.095 -0.026 0.009 0.021 0.006 -0.038 0.735
BY
I 0.001 -0.002 -0.002 0.032 0.040 -0.084 0.042 -0.038 0.006 0.004 0.148 0.559
TSD RESIDUAL EFFECT = 0.01 IRRIGATED RESIDUAL EFFECT = 0.02

DF - Days to 50 per cent flowering PH - Plant height (cm) PL - Panicle length (cm) FLL – Flag leaf length (cm),
FLW - Flag leaf Width (cm) FLA - Flag leaf area (cm2) GL –Grain Length (mm) GW – Grain Width (mm)
LB – Grain L: B ratio SI -100 seed weight (g) HI-Harvest index (%) BY -Biological yield (g/m2)
T – TSD condition I – Irrigated condition
Table 4.7: Correlation coefficient of grain yield and its component in without QTL lines
PH PL FLL FLW FLA GL GW LB SI HI BY GY

T 0.230 0.325 0.315 -0.177 0.064 0.044 -0.067 0.088 0.037 -0.189 -0.018 -0.191
DF
I 0.211 -0.090 0.105 -0.061 0.067 -0.062 -0.264 0.195 -0.114 -0.102 -0.031 -0.117

T 0.159 0.310 -0.031 0.181 0.053 -0.073 0.110 -0.061 -0.110 0.264 0.133
PH
I 0.502* *0.492 -0.096 0.327 0.223 0.115 0.058 -0.116 -0.102 0.259 0.099

T 0.074 0.014 -0.050 0.083 0.108 -0.005 0.190 0.029 -0.065 -0.037
PL
I 0.348 0.002 0.296 0.336 0.029 0.146 0.173 0.148 0.318 0.279

T -0.043 0.591** -0.077 0.135 -0.185 -0.306 0.002 0.059 -0.037


FLL
I -0.057 0.755** -0.043 0.141 -0.017 -0.074 -0.101 0.076 -0.021

T 0.765** -0.120 -0.099 -0.039 -0.102 0.176 -0.341 -0.149


FLW
I 0.596** 0.258 0.139 -0.040 0 .357 0.197 0.071 0.149

T -0.142 0.005 -0.142 -0.261 0.136 -0.251 -0.161


FLA
0.170 0.267 -0.081 0.235 0.052 0.130 0.100

T 0.275 0.617** 0.707** -0.140 0.208 0.103


GL
I 0.119 0.275 0.611** 0.183 0.207 0.241

T -0.581** 0.369 -0.067 0.357 0.226


GW
I -0.885** 0.171 0.034 0.168 0.140

T 0.296 -0.084 -0.107 -0.107


LB
I 0.073 0.003 -0.068 -0.040
T -0.101 0.206 0.130
SI
I 0.277 0.180 0.273

T -0.242 0.559*
HI
I 0.376 0.807**

T 0.650**
BY
I 0.835**

DF - Days to 50 per cent flowering PH - Plant height (cm) PL - Panicle length (cm) FLL – Flag leaf length (cm),
FLW - Flag leaf Width (cm) FLA - Flag leaf area (cm2) GL –Grain Length (mm) GW – Grain Width (mm)
LB – Grain L:B ratio SI -100 seed weight (g) HI-Harvest index (%) BY -Biological yield (g/m2)
GY - Grain yield (g/m2) T – TSD condition I – Irrigated condition

* Significant at 5% level of significance and ** Significant at 1% level of significance


Table 4.12: Path coefficient analysis showing direct and indirect effects of different yield contributing character on yield in without QTL
lines

DF PH PL FLL FLW FLA GL GW LB SI HI BY

T -0.002 0.010 0.003 -0.041 0.009 0.004 -0.003 -0.008 0.011 -0.001 -0.150 -0.015
DF
I -0.067 0.010 0.002 -0.028 0.012 0.020 0.003 -0.054 0.045 0.006 -0.058 -0.019

T -0.001 0.044 0.002 -0.040 0.002 0.012 -0.004 -0.008 0.014 0.001 -0.087 0.217
PH
I -0.014 0.003 -0.013 -0.132 0.019 0.098 -0.011 0.023 0.013 0.006 -0.057 0.163

T -0.001 0.007 0.011 -0.010 0.001 0.003 -0.006 0.012 -0.001 -0.004 0.023 -0.053
PL
I 0.006 0.001 -0.027 -0.093 0.001 0.089 -0.018 0.006 0.034 -0.009 0.083 0.200

T -0.001 0.014 0.001 -0.129 0.002 0.040 0.005 0.015 -0.023 0.007 0.002 0.049
FLL
I -0.007 0.001 -0.009 -0.268 0.011 0.226 0.002 0.029 -0.004 0.004 -0.057 0.048

T 0.000 -0.001 0.000 0.006 -0.056 0.052 0.009 -0.011 -0.005 0.002 0.140 -0.280
FLW
I 0.004 -0.001 -0.000 0.015 -0.197 0.179 -0.014 0.029 -0.009 -0.018 0.111 0.045

T 0.000 0.008 0.001 -0.076 -0.043 0.068 0.010 0.001 -0.018 0.006 0.108 -0.206
FLA
I -0.004 0.001 -0.008 -0.202 -0.117 0.300 -0.009 0.056 -0.019 -0.012 0.029 0.082

T 0.000 0.002 0.001 0.010 0.007 -0.010 -0.071 0.031 0.077 -0.016 -0.111 0.171
GL
I 0.004 0.001 -0.002 0.011 -0.050 0.051 -0.054 0.025 0.065 -0.031 0.103 0.130

T 0.000 -0.003 0.001 -0.017 0.006 0.000 -0.020 0.112 -0.073 -0.009 -0.054 0.293
GW
I 0.018 0.003 -0.001 -0.037 -0.027 0.080 -0.006 0.208 -0.209 -0.009 0.019 0.105

T 0.000 0.005 0.000 0.024 0.002 -0.010 -0.044 -0.065 0.125 -0.007 -0.066 -0.087
LB
I -0.013 0.001 -0.004 0.004 0.007 0.024 -0.015 -0.184 0.236 -0.004 0.002 -0.043

T 0.000 -0.003 0.002 0.039 0.006 -0.018 -0.050 0.041 0.037 -0.023 -0.080 0.169
SI
I 0.001 -0.001 -0.004 0.019 -0.070 0.070 -0.033 0.036 0.017 -0.051 0.156 0.113

T 0.001 -0.005 0.000 0.000 -0.010 0.009 0.010 -0.008 -0.011 0.002 0.795 -0.198
HI
I 0.007 -0.000 -0.004 0.026 -0.038 0.015 -0.010 0.007 0.001 -0.014 0.562 0.236

T 0.000 0.012 -0.001 -0.008 0.019 -0.017 -0.015 0.040 -0.013 -0.005 -0.192 0.820
BY
I 0.002 0.001 -0.008 -0.020 -0.014 0.039 -0.011 0.035 -0.016 -0.009 0.211 0.628

TSD RESIDUAL EFFECT = 0.01 IRRIGATED RESIDUAL EFFECT = 0.01


DF - Days to 50 per cent flowering PH - Plant height (cm) PL - Panicle length (cm) FLL – Flag leaf length (cm),
FLW - Flag leaf Width (cm) FLA - Flag leaf area (cm2) GL –Grain Length (mm) GW – Grain Width (mm)
LB – Grain L: B ratio SI -100 seed weight (g) HI-Harvest index (%) BY -Biological yield (g/m2)
T – TSD condition I – Irrigated condition

Table 4.8: Correlation coefficient of grain yield and its component in check lines

PH PL FLL FLW FLA GL GW LB SI HI BY GY

T -0.846 0.161 -0.664 0.369 -0.446 0.252 0.176 0.114 -0.501 -0.898* -0.651 -0.911*
DF
I -0.958* -0.090 -0.810 0.521 0.001 -0.203 -0.711 0.228 -0.548 -0.280 -0.395 -0.301

T -0.299 0.625 -0.181 0.548 -0.428 0.083 -0.376 0.385 0.669 0.414 0.672
PH
I 0.057 0.768 -0.658 -0.151 -0.016 0.694 -0.387 0.414 0.228 0.289 0.232

T -0.337 0.554 -0.046 0.566 0.133 0.334 -0.501 -0.898* -0.651 -0.911*
PL
I -0.107 0.147 0.023 0.621 -0.396 0.707 0.448 0.556 0.519 0.563

FLL T -0.344 0.803 -0.195 -0.226 -0.021 0.385 0.669 0.414 0.672

I -0.193 0.427 0.110 0.831 -0.399 0.243 0.088 0.196 0.073

FLW T 0.256 -0.076 0.666 -0.382 0.175 0.160 0.103 0.129

I 0.800 0.508 -0.203 0.478 -0.069 0.111 -0.030 0.016

FLA T -0.204 0.171 -0.220 0.415 0.523 0.522 0.555

I 0.517 0.346 0.162 0.072 0.165 0.086 0.062

GL T -0.351 0.876 0.519 0.165 0.198 0.135

I -0.070 0.801 0.737 0.547 0.608 0.563

GW T -0.751 -0.656 -0.334 -0.413 -0.347


I -0.644 0.176 0.232 0.169 0.151

LB T 0.657 0.248 0.337 0.237

I 0.457 0.242 0.329 0.300

SI T 0.756 0.523 0.726

I 0.667 0.835 0.769

HI T 0.724 0.994*

I 0.837 0.958*

BY T 0.791

I 0.548

DF - Days to 50 per cent flowering PH - Plant height (cm) PL - Panicle length (cm) FLL – Flag leaf length (cm),
FLW - Flag leaf Width (cm) FLA - Flag leaf area (cm2) GL –Grain Length (mm) GW – Grain Width (mm)
LB – Grain L: B ratio SI -100 seed weight (g) HI-Harvest index (%) BY -Biological yield (g/m2)
GY - Grain yield (g/m2) T – TSD condition I – Irrigated condition

* Significant at 5% level of significance and **Significant at 1% level of significance


Table 4.12: Path coefficient analysis showing direct and indirect effects of different yield contributing character on yield in Check
lines

DF PH PL FLL FLW FLA GL GW LB SI HI BW

T 0.114 0.065 0.068 0.380 -0.222 -0.274 -0.014 0.022 -0.049 -0.259 -0.410 -0.244
DF
I 1.957 1.038 0.328 -3.719 -1.364 -0.001 -0.710 3.259 0.048 0.486 -1.708 1.055
T -0.097 -0.076 -0.126 -0.358 0.109 0.337 0.024 0.010 0.161 0.199 0.306 0.155
PH
I -1.087 -1.084 -0.208 3.525 1.721 0.015 -0.056 -3.181 -0.081 -0.367 1.386 -0.770
T 0.018 0.023 0.423 0.193 -0.334 -0.028 0.032 -0.016 -0.143 0.090 0.073 0.039
PL
I -0.175 -0.061 -3.656 -0.491 -0.384 -0.002 2.177 1.815 0.148 -0.397 3.386 -1.384
T -0.076 -0.048 -0.142 -0.573 0.208 0.494 0.011 -0.028 0.009 0.215 0.239 0.195
FLL
I -1.586 -0.832 0.391 4.591 0.504 -0.044 0.384 -3.807 -0.084 -0.215 0.535 -0.523
T 0.042 0.014 0.234 0.197 -0.603 0.157 0.004 0.082 0.164 -0.241 -0.152 -0.076
FLW
I 1.020 0.713 -0.537 -0.884 -2.616 -0.083 1.783 0.932 0.100 0.061 0.676 0.079
T -0.051 -0.042 -0.019 -0.460 -0.154 0.615 0.011 0.021 0.094 0.081 0.152 0.152
FLA
I 0.001 0.164 -0.083 1.958 -2.092 -0.103 1.814 -1.586 0.034 -0.064 1.006 -0.230
T 0.029 0.033 0.240 0.112 0.046 -0.125 -0.056 -0.043 -0.375 0.269 0.075 0.074
GL
I -0.397 0.017 -2.269 0.505 -1.329 -0.054 3.510 0.319 0.169 -0.654 3.335 -1.621
T 0.020 -0.006 0.056 0.129 -0.402 0.105 0.020 0.124 0.321 -0.340 -0.153 -0.155
GW
I -1.392 -0.752 1.448 3.813 0.532 -0.036 -0.244 -4.584 -0.136 -0.157 1.414 -0.451
T 0.013 0.029 0.141 0.231 0.012 -0.136 -0.049 -0.093 -0.428 0.340 0.113 0.126
LB
I 0.447 0.419 -2.584 -1.833 -1.251 -0.017 2.812 2.952 0.211 -0.406 1.475 -0.877
T -0.057 -0.029 0.074 -0.238 0.281 0.096 -0.029 -0.081 -0.281 0.517 0.346 0.195
SI
I -1.073 -0.449 -1.637 1.115 0.180 -0.008 2.588 -0.808 0.096 -0.888 4.065 -2.229
T -0.103 -0.051 0.068 -0.300 0.200 0.205 -0.009 -0.041 -0.106 0.391 0.457 0.271
HI
I -0.549 -0.247 -2.032 0.403 -0.290 -0.017 1.921 -1.064 0.051 -0.592 6.094 -2.233
T -0.074 -0.032 0.044 -0.299 0.123 0.250 -0.011 -0.051 -0.144 0.270 0.331 0.374
BY
I -0.774 -0.313 -1.897 0.901 0.078 -0.009 2.132 -0.774 0.069 -0.741 5.099 -2.669
TSD RESIDUAL EFFECT = 0.01 IRRIGAT ED RESIDUAL EFFECT = 0.01

DF - Days to 50 per cent flowering PH - Plant height (cm) PL - Panicle length (cm) FLL – Flag leaf length (cm),
FLW - Flag leaf Width (cm) FLA - Flag leaf area (cm2) GL –Grain Length (mm) GW – Grain Width (mm)
LB – Grain L: B ratio SI -100 seed weight (g) HI-Harvest index (%) BY -Biological yield (g/m2)
T – TSD condition I – Irrigated condition

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