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Sustainable Development and Biodiversity 10

Vijay Rani Rajpal

S. Rama Rao
S.N. Raina Editors

Gene Pool
Diversity
and Crop
Improvement
Volume 1

123
Sustainable Development and Biodiversity

Volume 10

Series editor
Kishan Gopal Ramawat
Botany Department, M.L. Sukhadia University, Udaipur, Rajasthan, India
This book series provides complete, comprehensive and broad subject based reviews
about existing biodiversity of different habitats and conservation strategies in the
framework of different technologies, ecosystem diversity, and genetic diversity.
The ways by which these resources are used with sustainable management and
replenishment are also dealt with. The topics of interest include but are not restricted
only to sustainable development of various ecosystems and conservation of hotspots,
traditional methods and role of local people, threatened and endangered species,
global climate change and effect on biodiversity, invasive species, impact of various
activities on biodiversity, biodiversity conservation in sustaining livelihoods and
reducing poverty, and technologies available and required. The books in this series
will be useful to botanists, environmentalists, marine biologists, policy makers,
conservationists, and NGOs working for environment protection.

More information about this series at http://www.springer.com/series/11920

Vijay Rani Rajpal S. Rama Rao
•

S.N. Raina
Editors

Gene Pool Diversity

and Crop Improvement
Volume 1

123
Editors
Vijay Rani Rajpal S.N. Raina
Delhi University Amity University of Biotechnology
Delhi Noida, Uttar Pradesh
India India

S. Rama Rao
Department of Biotechnology
and Bioinformatics
North Eastern Hill University
Shillong, Megalaya
India

ISSN 2352-474X ISSN 2352-4758 (electronic)

Sustainable Development and Biodiversity
ISBN 978-3-319-27094-4 ISBN 978-3-319-27096-8 (eBook)
DOI 10.1007/978-3-319-27096-8

Library of Congress Control Number: 2015957776

© Springer International Publishing Switzerland 2016

This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part
of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations,
recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission
or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar
methodology now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this
publication does not imply, even in the absence of a specific statement, that such names are exempt from
the relevant protective laws and regulations and therefore free for general use.
The publisher, the authors and the editors are safe to assume that the advice and information in this
book are believed to be true and accurate at the date of publication. Neither the publisher nor the
authors or the editors give a warranty, express or implied, with respect to the material contained herein or
for any errors or omissions that may have been made.

Printed on acid-free paper

This Springer imprint is published by SpringerNature

The registered company is Springer International Publishing AG Switzerland
Preface

To respond to an ever-increasing need of food and fibre by the growing world’s

population, standing today at 7 billion and expected to reach 9 billion by 2050,
there is a pressing need to increase crop productivity. This can be achieved by
developing cultivars with better grain yield and high nutritive value through plant
breeding in traditional crop plants and in supplementary crops identified by
International Plant Genetic Resources Institute (IPGRI) and Consultative Group on
International Agriculture (CGIAR). The genetic improvement can also be achieved
by using biotechnological tools for alien gene transfer and engineering the traits in
plants that are otherwise difficult using conventional plant breeding approaches, by
newer methods to precisely and rapidly screen for traits of interest in the progeny,
and by cytogenetic manipulations. Knowledge of genetic diversity, locked in the
germplasm resources of the crop plants and wild relatives constituting primary,
secondary, and tertiary gene pools, and the genome(s) characterization is essential
for crop improvement and developing gene transfer strategies by multitude of tools
that are now available.
This book addresses aforementioned issues in several crop species. Each chapter
elucidates an authoritative account on the topic. We are sincerely grateful to all the
authors for their valuable contributions. We would like to acknowledge coopera-
tion, patience, and support of our contributors, who have put in their serious efforts
to ensure a high scientific quality of this book with up-to-date information. We
thank Dr. K.G. Ramawat for motivating us to take up this assignment. Sincere
thanks are due to Khushboo Arora for her help during the editing process. This
work could not be completed without the active support of Springer team who took
pains in streamlining the production process. We particularly appreciate Dr. Valeria
for her continued support. Vijay Rani Rajpal is sincerely grateful to her husband
Susheel Rajpal and daughter Navya Rajpal for their patience and support during the
entire period of this book project.

v
vi Preface

Plant breeders, taxonomists, geneticists, cytogeneticists, molecular biologists,

and biotechnologists will greatly benefit from this book. We sincerely hope that this
book will serve as a milestone towards achieving meaningful plant genetic
improvement to meet the ever-increasing requirements of food and fibre of this
world.

Vijay Rani Rajpal

S. Rama Rao
S.N. Raina
Contents

1 Leymus racemosus: A Potential Species of Gene Pool Enrichment

for Wheat Improvement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Yasir Serag Alnor Gorafi and Hisashi Tsujimoto
2 Arachis Gene Pools and Genetic Improvement in Groundnut . . . . . 17
Anurudh K. Singh and S.N. Nigam
3 Genetic Resources of Chickpea (Cicer arietinum L.)
and Their Utilization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Deepak Ohri
4 Genetic Improvement of Cotton . . . . . . . . . . . . . . . . . . . . . . . . . . 105
S. Manickam and A.H. Prakash
5 Genetic Diversity and Germplasm Patterns in Brassica juncea . . . . 163
S.S. Banga and Shashi Banga
6 Potato Diversity and Its Genetic Enhancement . . . . . . . . . . . . . . . 187
Avinash Srivastava, Vinay Bhardwaj, BP Singh
and SM Paul Khurana
7 Genome Plasticity in Buckwheat . . . . . . . . . . . . . . . . . . . . . . . . . . 227
Nikhil K. Chrungoo, Lashaihun Dohtdong and Upasna Chettry
8 Origin of Genetic Variability and Improvement
of Quinoa (Chenopodium quinoa Willd.) . . . . . . . . . . . . . . . . . . . . 241
Atul Bhargava and Deepak Ohri
9 Emerging Invaders from the Cultivated Croplands:
An Invasion Perspective . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271
Neha Goyal and Gyan Prakash Sharma
10 Chromosome Engineering for High Precision Crop Improvement . . . 291
Harinder Kumar Chaudhary, Vineeta Kaila, Shoukat Ahmad Rather,
Navdeep Singh Jamwal and Anila Badiyal

vii
viii Contents

11 Overview of Progress and Potentials of Improving Commonly

Used Allium species in India . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 325
R.N. Gohil and Veenu Kaul
12 Utilization of Germplasm for the Genetic Improvement of Mung
bean [Vigna radiata (L.) Wilczek]: The Constraints
and the Opportunities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 367
Ruchi Vir, Suman Lakhanpaul, Sonal Malik, Sooraj Umdale
and Kangila Venkataramana Bhat
13 Genetic Improvement in the Genus Eleusine . . . . . . . . . . . . . . . . . 393
Renuka Agrawal and Ankur Maheshwari
14 An Ancient Medicinal Plant at the Crossroads of Modern
Horticulture and Genetics: Genetic Resources and Biotechnology
of Sea Buckthorn (Hippophae L., Elaeagnaceae) . . . . . . . . . . . . . . 415
Igor V. Bartish
15 Unlocking the Potential of Genetic Resources for Improvement
of Sesame (Sesamum indicum L.): The Current Scenario . . . . . . . . 447
Vibhuti Singh, Sachin Kumar, Amrita Singh, Niti Pathak Bhaduri,
Kangila Venkataramana Bhat and Suman Lakhanpaul

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 481
Contributors

Renuka Agrawal Department of Botany, Miranda House, University of Delhi,

Delhi, India
Anila Badiyal Molecular Cytogenetics and Tissue Culture Laboratory,
Department of Crop Improvement, CSK Himachal Pradesh Agricultural University,
Palampur, India
S.S. Banga Department of Plant Breeding and Genetics, Punjab Agricultural
University, Ludhiana, India
Shashi Banga Department of Plant Breeding and Genetics, Punjab Agricultural
University, Ludhiana, India
Igor V. Bartish Department of Genetic Ecology, Institute of Botany, Academy of
Sciences of Czech Republic, Pruhonice, Czech Republic
Niti Pathak Bhaduri Department of Botany, University of Delhi, Delhi, India
Vinay Bhardwaj Central Potato Institute Shimla, Shimla, HP, India
Atul Bhargava Amity University Uttar Pradesh (Lucknow Campus), Lucknow,
Uttar Pradesh, India
Kangila Venkataramana Bhat National Bureau of Plant Genetic Resources, Pusa
Campus, New Delhi, India
Harinder Kumar Chaudhary Molecular Cytogenetics and Tissue Culture
Laboratory, Department of Crop Improvement, CSK Himachal Pradesh
Agricultural University, Palampur, India
Upasna Chettry Plant Molecular Biology Laboratory, UGC-Centre for Advanced
Studies in Botany, North-Eastern Hill University, Shillong, India
Nikhil K. Chrungoo Plant Molecular Biology Laboratory, UGC-Centre for
Advanced Studies in Botany, North-Eastern Hill University, Shillong, India

ix
x Contributors

Lashaihun Dohtdong Plant Molecular Biology Laboratory, UGC-Centre for

Advanced Studies in Botany, North-Eastern Hill University, Shillong, India
R.N. Gohil Department of Botany, University of Jammu, Jammu, India
Yasir Serag Alnor Gorafi Arid Land Research Center, Tottori University, Tottori,
Japan; Agricultural Research Corporation, Wad Medani, Sudan
Neha Goyal Department of Environmental Studies, University of Delhi, Delhi,
India
Navdeep Singh Jamwal Molecular Cytogenetics and Tissue Culture Laboratory,
Department of Crop Improvement, CSK Himachal Pradesh Agricultural University,
Palampur, India
Vineeta Kaila Molecular Cytogenetics and Tissue Culture Laboratory,
Department of Crop Improvement, CSK Himachal Pradesh Agricultural University,
Palampur, India
Veenu Kaul Department of Botany, University of Jammu, Jammu, India
SM Paul Khurana Faculty of Science, Engineering & Technology, University
Science Instrumentation Centre, Amity University, Gurgaon, Haryana, India
Sachin Kumar Department of Botany, University of Delhi, Delhi, India
Suman Lakhanpaul Department of Botany, University of Delhi, Delhi, India
Ankur Maheshwari Department of Zoology, Zakir Husian Delhi College,
University of Delhi, New Delhi, India
Sonal Malik Department of Botany, University of Delhi, Delhi, India
S. Manickam ICAR-Central Institute for Cotton Research, Regional Station,
Coimbatore, India
S.N. Nigam Hyderabad, Telangana, India
Deepak Ohri Amity University Uttar Pradesh (Lucknow Campus), Lucknow, UP,
India
A.H. Prakash ICAR-Central Institute for Cotton Research, Regional Station,
Coimbatore, India
Shoukat Ahmad Rather Molecular Cytogenetics and Tissue Culture Laboratory,
Department of Crop Improvement, CSK Himachal Pradesh Agricultural University,
Palampur, India
Gyan Prakash Sharma Department of Environmental Studies, University of
Delhi, Delhi, India
Amrita Singh Department of Botany, University of Delhi, Delhi, India
Anurudh K. Singh Gurgaon, Haryana, India
Contributors xi

BP Singh Central Potato Institute Shimla, Shimla, HP, India

Vibhuti Singh Department of Botany, University of Delhi, Delhi, India
Avinash Srivastava Central Potato Research Station, Shillong, Meghalaya, India
Hisashi Tsujimoto Arid Land Research Center, Tottori University, Tottori, Japan
Sooraj Umdale National Bureau of Plant Genetic Resources, Pusa Campus, New
Delhi, India
Ruchi Vir Department of Botany, University of Delhi, Delhi, India
Chapter 1
Leymus racemosus: A Potential Species
of Gene Pool Enrichment for Wheat
Improvement

Yasir Serag Alnor Gorafi and Hisashi Tsujimoto

Abstract Leymus racemosus is a wild species belonging to tribe Triticeae

(Poaceae), which includes important cereal crops such as bread wheat and barley.
This perennial species grows along the coast and in dry lands, and is reportedly
tolerant to various biotic and abiotic stresses. Although L. racemosus is evolu-
tionarily distant from wheat (Triticum spp.) within the tribe, it has been successfully
hybridized with wheat, and several wheat–L. racemosus chromosome introgression
lines have been selected from among the backcrossed progenies of the hybrid. L.
racemosus is, therefore, a promising wild species for wheat improvement. The
production of wheat, as one of the world’s most important staple cereals, must
increase to provide food for growing population, but wheat is threatened by the
effects of climate change, especially increased drought and heat. In addition,
depletion of natural resources makes the likelihood of success for increasing future
productivity unclear. Maximizing yield to help achieve food security under such
challenging conditions will not be easy. Wheat productivity could be improved by
enhancing tolerance to biotic and abiotic stress, and tolerance to soil macro- and
micronutrient deficiencies or toxicities. However, the genetic base of variation
available for most of these traits is very narrow in the available elite germplasm.
Wild relatives of wheat, including L. racemosus, are an important source of wheat
genetic variation and have contributed much to the improvement of wheat pro-
ductivity. This review describes the potential of this wild wheat relative as a
germplasm for wheat improvement, and the characteristics that affect its efficient
use in breeding, including its chromosomes, traits, and some genes that have been
identified and transferred from this species to common wheat.

Keywords Leymus Wheat Germplasm enhancement Drought Chromosome


introgression line

Y.S.A. Gorafi
H. Tsujimoto (&)

Arid Land Research Center, Tottori University, 1390 Hamasaka, Tottori 680-0001, Japan
e-mail: [email protected]
Y.S.A. Gorafi
Agricultural Research Corporation, PO Box 126, Wad Medani, Sudan

© Springer International Publishing Switzerland 2016 1

V.R. Rajpal et al. (eds.), Gene Pool Diversity and Crop Improvement,
Sustainable Development and Biodiversity 10,
DOI 10.1007/978-3-319-27096-8_1
2 Y.S.A. Gorafi and H. Tsujimoto

1.1 Introduction

Wheat (Triticum spp.) is the world’s most widely grown grain and provides nearly
20 % of the calories humans require (FAOstat 2007). The world’s population is
growing exponentially and will exceed nine billion by 2050. According to recent
FAO estimates, projected demand will require global agricultural production to
increase by about 60 % relative to yields obtained between 2005 and 2007.
Between 2010 and 2012, nearly one in eight people in the world (870 million) had
less than the minimum amount of food required. Of these, 852 million comprised up
to 15 % of the total population of developing countries (Ogbonnaya et al. 2013).
Further, impact of climate change is expected to reduce wheat gain yield (Yang
et al. 2013; Lobell et al. 2011), which will further complicate food production in the
future. The requirement for higher productivity under more limiting conditions,
whether by increasing yield or acreage, is an urgent objective that must be met.
However, the enhancement of yield is limited by biotic and abiotic stresses and
limited availability of arable land.
The modern bread wheat (Triticum aestivum) , which accounts for 95 % of the
production of all wheat species, has evolved only 8500 years ago from a rare
interspecific hybridization event between tetraploid and diploid wheat (Kihara
1944, 1966; McFadden and Sears 1944; Qi et al. 2007). Because this event is
thought to have occurred only once (Curtis and Halford 2014), all modern wheat is
thought to have descended from that single cross. Consequently, the genetic vari-
ability of hexaploid bread wheat germplasm has a rather narrow base, and after
domestication and several decades of extensive selection and breeding, this already
limited genetic base has become even narrower (Reif et al. 2005). With such a
narrow base, the required increases in wheat yields are hard to imagine. Therefore,
there is now growing interest in the great genetic diversity present in wild relatives
of wheat. Certain traits in some of these wild relatives have already been charac-
terized and successfully used to improve the tolerance of wheat to abiotic (such as
heat, salinity, and drought) and biotic stresses (e.g., diseases, insects, and
kit-nematodes), and to improve nutrient use efficiency, grain yield, and bread
making quality (Eastwood et al. 1994; Jiang et al. 1994; Cox et al. 1995; Gatford
et al. 2002; Marais et al. 1994; Martín-Sánchez et al. 2003; Dreccer et al. 2004;
Wang et al. 2010; Garg et al. 2009; Liu et al. 2013), via molecular biology and
cytogenetic techniques.
This review sheds light on the potential of an important wild relative of wheat, L.
racemosus for wheat improvement, and describes several characteristics; including
its chromosomes, useful traits, and genes identified to date that could be transferred
to improve common wheat.
1 Leymus racemosus: A Potential Species … 3

1.2 Botanical Information for Leymus racemosus

Among the wild relatives of wheat, the genus Leymus (wildrye) is a distantly related
source of many potentially adaptive alien genes that could be useful for the
breeding of common wheat. Leymus is an allopolyploid genus with chromosomes
numbers ranging from 2n = 4× = 28 to 2n = 12× = 84 in the tribe Triticeae, which
comprises about 50 species and subspecies worldwide (Löve 1984). The genome of
tetraploid L. racemosus is comprised of two genomes of different origins (2n = 28,
NsNsXmXm); the Ns genome originated from Psathyrostachys, while the origin of
the Xm genome remains unknown (Zhang and Dvorak 1991; Wang and Jensen
1994; Wang et al. 1994, 2006).
L. racemosus (Lam.) Tzvelev (Fig. 1.1a), is a perennial grass that is also known
by the common name ‘mammoth wildrye’ as well as other names including Elymus
arenarius var. giganteus, E. giganteus, E. racemosus, and L. giganteus (PLANTS
Database 2010; http://plants.usda.gov/). The species L. racemosus distributes in a
range of dry coastal or inland sites from Central Asia to Eastern Europe (Kishii
et al. 2004). It is also found growing in a variety of environments including saline,
alkaline, dry, or semi-dry lands and forests (Fan et al. 2009). Different from the
other species, in Leymus only L. racemosus and L. mollis have exceptionally large
spikes, strong rhizomes, and vigorous growth (Fig. 1.1b).
L. racemosus has been used for ecological purposes such as stabilizing and
protecting sand dunes from wind erosion, revegetating mine tailings, and making
firebreaks and green strips. It is also used to create wildlife habitat for birds (St.
John 2010). Monsen et al. (2004) reported that L. racemosus is generally unsuitable
and unpalatable for grazing animal.

(a) (b)

(c)

Fig. 1.1 Morphology of Leymus racemosus and its addition lines. a L. racemosus maintained in a
pot of 50 cm diameter. This plant is perennial, stress-tolerant and forms long rhizomes as indicated
by the arrow. b The long rhizome of L. mollis, the species most closely resembling L. racemosus.
c Spikes of wheat–L. racemosus chromosome introgression lines (CILs). The morphology of the
CILs is similar to common wheat
4 Y.S.A. Gorafi and H. Tsujimoto

L. racemosus is quite salt- and drought-tolerant (McGuire and Dvorak 1981) and
is also resistant to various diseases including scab (Mujeeb-Kazi et al. 1983). Our
observations of L. racemosus plants growing at the Arid Land Research Center,
Tottori, Japan (35°32′N, 134°13′E), confirmed the ability of this species to tolerate
heat stress; plants grew vigorously and flowered from May to June, and
grain-filling, which is particularly a heat-sensitive stage, succeeded from July
through August during high summer temperatures and humidity.

1.3 Crossability of Leymus racemosus with Wheat

and Chromosome Introgression Lines

Several Leymus species, including L. racemosus, have been successfully hybridized

and backcrossed with wheat. The first pentaploid hybrid between wheat and L.
racemosus, which included the genomes ABDNsXm was developed by
Mujeeb-Kazi and Rodriguez (1981). The second attempt of hybridization was made
in China, aimed at introducing the genes for scab resistance from L. racemosus into
the common wheat. Wang et al. (1986, 1991) characterized the cytogenetics of the
F1 hybrid between wheat and L. racemosus and its BC1 and BC2 derivatives.
Subsequently, seven disomic addition lines and two di-telosomic addition lines
were developed and screened for scab resistance (Chen et al. 1993, 1995; Ren et al.
1996; Sun et al. 1997, 1998). Lu et al. (1995) developed one disomic addition line
and one double-disomic addition–substitution line via anther culture. Another five
disomic addition lines and several additional double-disomic addition, di-telosomic
addition, and disomic substitution lines were characterized in 23 lines derived from
the bread wheat cultivar ‘Chinese Spring’ (CS) (Qi et al. 1997). Kishii et al. (2004)
produced ten chromosome addition lines using a L. racemosus accession collected
along the Bulgarian Black Sea coast (Fig. 1.1c). Table 1.1 summarizes the wheat–L.
racemosus chromosome introgression lines maintained in the Tottori Alien
Chromosome Bank for Wheat (TACBOW) that is funded by the National
Bio-resources project (NBRP-wheat), and the chromosome information for each.

1.4 Use of Genes from Leymus racemosus for Wheat

Improvement

There are two possible strategies currently available to use genes from L. racemosus,
or other wild relatives of wheat, for wheat improvement. In the first approach, the
desired phenotype of wild species is identified by direct examination of the species.
This is followed by crossing wheat and the wild species to produce a sterile F1 hybrid
by embryo rescue. Then, the F1 plants are repeatedly backcrossed to wheat to
generate CILs (including addition, substitution, and translocation lines) that harbor
1 Leymus racemosus: A Potential Species … 5

Table 1.1 List of the wheat–Leymus chromosome introgression lines produced or maintained at
the Tottori Alien Chromosome Bank of Wheat supported by NBRP-wheat
Strain ID Strain name and Homoeologous Number of References
chromosome name group chromosomes
TACBOW0001a Leymus racemosus 2 44 Kishii et al.
A addition (2004)
TACBOW0003 L. racemosus E NDb 44 Kishii et al.
addition (2004)
TACBOW0004 L. racemosus F 4 44 Kishii et al.
addition (2004)
TACBOW0005 L. racemosus H 3 44 Kishii et al.
addition (2004)
TACBOW0006 L. racemosus I 5 44 Kishii et al.
addition (2004)
TACBOW0008 L. racemosus k 6 44 Kishii et al.
addition (2004)
TACBOW0009 L. racemosus l 2 44 Kishii et al.
addition (2004)
TACBOW0010 L. racemosus n 3,7 44 Kishii et al.
addition (2004)
TACBOW0011 L. racemosus H 3 42 Kishii et al.
substitution (2004)
TACBOW0012 L. racemosus 2 44 Qi et al.
(TA#7643)a 2Lr#1 addition (1997)
TACBOW0013 L. racemosus 5 44 Qi et al.
(TA#7646) 5Lr#1 addition (1997)
TACBOW0014 L. racemosus 6 44 Qi et al.
(TA#7648-1) 7Lr#1 addition (1997)
TACBOW0015 L. racemosus 3, 7 44 Qi et al.
(TA#7648-3) 7Lr#1 addition (1997)
TACBOW0016 L. racemosus ? NAc 44 Qi et al.
(TA#7652) Lr#1 addition (1997)
TACBOW0017 L. racemosus 2 42 Qi et al.
(TA#7644) 2Lr#1 substitution (1997)
a
TA#7643: Number from the Kansas State University gene bank
b
ND: not determined
c
NA: not available

entire or partial chromosomes from the wild species. Finally, these CILs are screened
to identify the line that has the desired phenotype. The advantage of this method is
reliability; and because useful traits can be easily followed from the wild species to
common wheat and it may be particularly efficient for qualitative traits. However,
this strategy is time-consuming, and expected traits from the wild species may not
exhibit sufficient expression in the genetic background of common wheat. This
happened especially in case of quantitative characters. In addition, wild relatives
such as L. racemosus are often perennials with substantial vegetative biomass, small
seeds, and poor agronomic traits. This nature hinders proper evaluation of their traits
6 Y.S.A. Gorafi and H. Tsujimoto

and response to different environmental conditions. Thus, it is not an easy task to

hunt for the useful genes or traits from wild species.
The second strategy for using genes from L. racemosus or other wild relatives of
wheat depends on phenotyping the CILs and not the wild species itself. The unique
property of these CILs is that they are phenotypically similar to wheat (Fig. 1.1c),
which eases evaluation for any useful genes they might carry. The advantage of this
method is that the presence of these genes in the wheat genetic background allows
evaluation of their positive or negative impact results from interactions with the
wheat genome, and the influence of the wheat genome on the expression of the
alien genes. Once a CIL with desired phenotype is identified, translocation lines
could be developed for a target chromosome that bears the desired phenotype.
Translocation lines could then be screened to identify lines with the desired
phenotype.
The utility of CILs is not limited to wheat breeding; they can also be used for
other applications. For example, Kikuchi et al. (2009) and Cho et al. (2011a) used
wheat–L. racemosus CILs to investigate the effects of heavy-ion beams and the
DNA methylation inhibitor zebularine on chromosome structure and rearrange-
ment. As those CILs harbor a pair of alien chromosomes that serve as markers that
can be easily detected by genomic in situ hybridization (GISH), Cho et al. (2011b)
used double monosomic addition lines carrying L. racemosus and L. mollis chro-
mosomes to study meiotic chromosome pairing. The line having two homologous
chromosome painted in different colors by GISH was quite useful for investigating
factors affecting meiotic chromosome pairing and recombination.

1.5 Impact of Leymus Genes on Improvement

of Wheat Traits

Within the last few decades several wheat–L. racemosus CILs have been developed
and evaluated, and useful chromosomes and traits have been determined, and
analyzed (Chen et al. 2005; Qi et al. 2008; Wang and Chen 2008; Subbarao et al.
2007; Mohammed et al. 2013, 2014). These lines include useful genes for disease
resistance, biological nitrification inhibition, aluminum tolerance, and heat stress
tolerance.

1.5.1 Scab Resistance

Mujeeb-Kazi et al. (1983) reported that L. racemosus is highly resistant to scab,

also known as Fusarium head blight. A wheat breeding program was later initiated
in China to transfer this scab tolerance to common wheat. Three scab-resistant
wheat–L. racemosus disomic addition lines (DALr.2, DALr.7, and DALr.14) were
1 Leymus racemosus: A Potential Species … 7

identified (Chen et al. 1993, 1995). Qi et al. (1997) reported that the additional
chromosomes in DALr.2 and DALr.14 belong to homoeologous group 7 and 5,
respectively, so they were designated 7Lr#1 and 5Lr#1. From these disomic
addition lines, new wheat–L. racemosus translocation lines were then developed by
inducing chromosome breakage with irradiation and gametocidal genes (Chen et al.
2005). In that study, nine scab-resistant wheat–L. racemosus translocation lines
were identified. In one of these lines, NAU614, the long arm of 5Lr#1 had been
translocated to wheat chromosome 6B. In four other lines, NAU601, NAU615,
NAU617, and NAU635, part of the short arm of 7Lr#1 had been transferred to
various wheat chromosomes. Another four lines, NAU611, NAU634, NAU633,
and NAU618 contained translocations involving Leymus chromosome Lr.7 and
various wheat chromosomes. The translocation lines containing a single alien
chromosome segment were more resistant to scab than was the susceptible wheat
parent CS but were less resistant than the resistant parent L. racemosus. At least
three resistance genes were identified in L. racemosus, including one located on
chromosome Lr.7, another that could be assigned to the long arm of 5Lr#1, and a
third that could be assigned to the short arm of 7Lr#1. One novel resistance gene
designated Fhb3 was also mapped to the distal region of the short arm of chro-
mosome 7Lr#1, and specific PCR-based markers were also developed to facilitate
marker-assisted selection in scab resistance breeding programs (Qi et al. 2008).

1.5.2 Biological Nitrification Inhibition

Biological nitrification is a serious problem in agriculture that results in costly

nitrogen loss. Subbarao et al. (2007) reported the first example of biological
nitrification inhibition (BNI) identified among the wild relatives of cereals in L.
racemosus, then introduced and successfully expressed this BNI trait in common
wheat. L. racemosus releases 20-fold higher amounts of BNI compounds than does
cultivated wheat. In their study, root exudates of cultivated wheat were unable to
inhibit biological nitrification in the soil, but the root exudates of L. racemosus
could suppress NO3− formation and maintain over 90 % of the inorganic nitrogen in
the soil as NH4+ for 60 days. Chromosomes Lr#n, Lr#J, and Lr#I were associated
with the high BNI capacity of L. racemosus (Fig. 1.2; Subbaro et al. 2007).
The BNI trait from the Lr#n chromosome was introduced into wheat by inducing a
Robertsonian translocation between wheat and the Lr#n chromosome (Kishii et al.
2008). Recently, a program was established at CIMMYT to identify and map the
genes responsible for BNI and introduce them into all commercial wheat cultivars
in order to reduce production costs and minimize the nitrogen pollution generated
by wheat production systems.
8 Y.S.A. Gorafi and H. Tsujimoto

Fig. 1.2 BNI release and dry 30

matter production of some
wheat–L. racemosus disomic 25
BNI released
addition lines (Subbarao et al. 20 (ATU/g root dry
2007)
weight/ day)
15
10
5
0

1.5.3 Aluminum Tolerance

Aluminum toxicity is a key factor that limits wheat production in the acidic soil
with pH below 5.5 (Evans and Kamprath 1970). Mohammed et al. (2013) screened
15 wheat–L. racemosus CILs for aluminum (Al) toxicity tolerance. L. racemosus
chromosomes Lr#A and Lr#E were found to significantly enhance relative root
growth (Fig. 1.3a, b; Mohammed et al. 2013) under Al stress and thereby increase
the Al tolerance of wheat. A line containing Lr#E chromosome was more
aluminum-tolerant than a line containing Lr#A chromosome at the highest Al
concentration tested (200 μM) (Mohammed et al. 2013). The introgressed chro-
mosomes did not affect the aluminum uptake or expression of the wheat ALMT1
gene that controls the exclusion of Al from root tips, but a gene or genes on
chromosome Lr#E did improve wheat cell membrane integrity of wheat under Al
stress conditions (Fig. 1.3c; Mohammed et al. 2013). The results suggest that there
are Al tolerance mechanisms other than exclusion of Al from the root tips.
Eventually, chromosome engineering using these two lines could result in wheat
cultivars with enhanced Al tolerance.

1.5.4 Heat Stress

Mohammed et al. (2014) examined the response of 14 wheat–L. racemosus CILs to

high temperature to determine whether they have potential for breeding improved
wheat cultivars. These introgression lines and their parent CS were evaluated either
in a growth chamber at the seedling stage and in the field at the reproductive stage
in two heat-stressed environments in Sudan. To be sure that the plants would be
exposed to heat stress at the reproductive stage; optimum and late planting strate-
gies were used. For seedlings under growth chamber conditions, the presence of
chromosomes Lr#A, Lr#I, Lr#l, and 2Lr#1 were found to improve heat stress
tolerance as measured by improved photosynthesis and mitochondrial electron
transport (Fig. 1.4a, b; Mohammed et al. 2014). Under field conditions, the
1 Leymus racemosus: A Potential Species … 9

Fig. 1.3 Performance of (a)

common wheat cultivar 120

Relative root growth (RRG) %

Chinese Spring (CS) and 94
100
Leymus racemosus 77
chromosome addition lines 80 66 66
57 53 53 53 55
under aluminum (Al) toxicity 60 47 49 50
38 42 44
conditions. a Relative root
40
growth (RRG) of CS and
addition lines grown under Al 20
toxicity conditions. Dark bars 0
indicate the Al tolerant lines.

CS
Lr#A
Lr#E
Lr#F
Lr#H
Lr#I
Lr#K
Lr#L
Lr#N
Lr#Hs
5Lr#1
7Lr#1
7Lr#1
?Lr#1
2Lr#1
b Sensitivity of CS and
tolerant lines Lr#A and Lr# E
to Al toxicity. c Plasma (b) CS Lr#A Lr#E
membrane integrity of CS and - AL +AL -AL +AL -AL +AL
tolerant lines Lr#A and Lr#E
under Al toxicity conditions
measured as percentage of
electrolyte leakage, dark bars
indicate the control and the
white bars indicate the Al
treatment (Mohammed et al.
2013)

(c)
80
Plasma membrane integrity (%)

65 67 64
61
57
60 50

40

20

0
CS Lr#A Lr#E

presence of chromosomes Lr#I and Lr#n enhanced grain number per spike and heat
tolerance (Table 1.2; Mohammed et al. 2014). Their results also indicated that
chromosome 7Lr#1 might improve yield potential as it has higher grain yield than
the wheat parent CS under non-stressed conditions (Table 1.2, Mohammed et al.
2014). Based on a heat susceptibility index (HSI) calculated from grain yield
obtained from optimum and late sowings, L. racemosus chromosomes Lr#I and
2Lr#1 were classified as highly heat tolerant, whereas Lr#A and 5Lr#1 were
classified as moderately heat tolerant (Table 1.2; Mohammed et al. 2014). This
study clearly indicated that several Leymus chromosomes have good potential for
improving wheat adaptation and tolerance to heat stress (Mohammed et al. 2014).
In addition to scab resistance, BNI, aluminum tolerance, and heat stress toler-
ance, lines carrying the Lr#J and Lr#l chromosomes also expressed gene(s) for
10 Y.S.A. Gorafi and H. Tsujimoto

Fig. 1.4 Reduction in (a)

chlorophyll fluorescence

Reduction of chlorophyll fluorescence

(a) and triphenyl tetrazolium 80
64
chloride (TTC) reduction 59 59
assay values (b) in CS and L. 60 53
racemosus addition lines 40 29
34 34
grown in a growth chamber 22 19 20 19 14
16 18
under normal and 20
heat-stressed conditions; dark
0
bars indicate lines with
significant differences from -20 -13
CS (Mohammed et al. 2014)

CS
Lr#A
Lr#E
Lr#F
Lr#H
Lr#I
Lr#K
Lr#l
Lr#n
Lr#Hs
2Lr#1
5Lr#1
7Lr#1
7Lr#1
2Lr#1
(b)
60
50
50
TTC value

40 37
33
27
30
21
17 16
20 13 15
11 10 11 13 12
10 7

0
CS
Lr…
Lr…
Lr#F
Lr…
Lr#I
Lr…
Lr#l
Lr#n
Lr…
2…
5…
7…
7…
2…
Table 1.2 Kernel number per spike, grain yield, and heat susceptibility index (HSI) of some
addition lines and their parent cultivar ‘Chinese Spring’ (CS) during evaluation for heat stress
tolerance at optimum planting (OP) and late planting (LP) in Shambat (2011/2012) and Gezira
(2012/2013), Sudan
Kernel number/spike Grain yield HSI
Shambat Gezira Shambat Gezira (g/m2)
(g/plant)
Line OP LP OP LP OP LP OP LP
Lr#A 53b 50b 49a 32a 1.9a 1.7a 122a 56a 0.59
Lr#I 68a 58a 47a 29b 2.3a 1.7a 132a 78a 0.24
a a
Lr#n 59 58 45b 26b 2.1a
1.9a 133a 50a 0.71
b b
2Lr#1 54 49 36b 27b 2.1a
1.7a 121a 74a 0.25
c c
5Lr#1 43 43 32b 14c 1.9a
1.8a 110a 56a 0.59
c b
7Lr#1 49 57 47a 36a 2.1a
1.9a 265a 29a 1.12
CS 55b 51b 37b 26b 1.5b 1.3b 51b 14b 1.40
a, b
Significant difference from CS
c
Compatible to CS

resistance to leaf or stem rust, and the Lr#k line possessed a gene for a novel seed
storage protein (Kishii 2011). Further, the Lr#H chromosome addition line exhib-
ited early heading (Kishii et al. 2004).
1 Leymus racemosus: A Potential Species … 11

1.6 Genetic Maps and Molecular Markers

Genetic maps and genomic information are essential for modern breeding programs
as they facilitate the assessment of genetic diversity for germplasm enhancement,
and ease the transfer of desired genetic traits by marker-assisted breeding
(Pérez-de-Castro 2012). Linkage mapping has been used extensively to identify
genes or QTLs in plant genomes that control important traits. It relies on random
recombination at meiosis and segregation of the recombinant chromosomes among
offsprings (Dear 2001). For genes introduced on alien chromosomes, it is difficult to
perform genetic analysis and construct genetic maps using CILs because no
recombination occurs at meiosis due to absence of homology between the alien
chromosome and the wheat chromosomes. Therefore, deletion mapping is the best
way to identify and map the genes located on the alien chromosome to allow their
transfer into wheat.
To create an efficient and informative deletion map, specific molecular markers
for the target chromosome are essential. No molecular markers specific for L.
racemosus have been developed to date. However, expressed sequence tags
(EST) markers from barley have been used to develop molecular markers that are
polymorphic between wheat and some important wild species (Hagras et al. 2005).
About 182 markers polymorphic between wheat and L. racemosus were identified
in their study. Although genetic maps are not yet available for L. racemosus, several
genetic maps have been constructed for other Leymus species. Molecular genetic
maps were constructed for two full-sib families (TTC1 and TTC2) derived from L.
cinereus and L. triticoides (Wu et al. 2003). The genetic maps of the TTC1 and
TTC2 families included a combined total of 1583 AFLP markers and 67 heterol-
ogous anchor markers (from other grass species) in 14 linkage groups. Two sets of
seven homoeologous linkage groups were tentatively identified based on synteny of
the heterologous anchor markers between wheat (Triticum spp.), barley (Hordeum
vulgare), and cereal rye (Secale cereale) (Wu et al. 2003; Larson et al. 2006). These
populations were found to segregate for genes and QTLs controlling growth habit
(Larson et al. 2006), forage quality (Larson and Mayland 2007), and seed shattering
(Larson and Kellog 2009).
From a collection of 28786 ESTs derived from subterranean rhizome and tiller
buds of L. triticoides × L. cinereus hybrids, 11281 unigene contigs were assembled
(Bushman et al. 2008). Of these, 9389 had at least one match with the rice genome.
In that study, 1798 SSR primers were designed of which 1575 were polymorphic
(Bushman et al. 2008). Further, Leymus EST-SSR primers that aligned to rice
chromosome 2 were tested for synteny in Leymus (Bushman et al. 2008; Kaur et al.
2008).
Using the same mapping populations, Larson et al. (2012) developed an inte-
grated consensus map including 375 AFLP, 350 SSR, and 48 heterologous markers
that cover 2381 cM. A total of 146 of these markers were tested in 17 wheat–L.
racemosus CILs and were found to be polymorphic between wheat and L. race-
mosus. All CILs tested positively for between 8 and 24 Leymus EST-SSR marker
12 Y.S.A. Gorafi and H. Tsujimoto

loci representing at least one homoeologous group; most of these matched the
previous chromosome identification (Larson et al. 2012).
Using suppressive-subtractive hybridization, Habora et al. (2012) identified 112
osmotic-stress-responsive genes (ESTs) from L. mollis that are differentially
expressed under osmotic stress. All these types of genetic information that are
available could be used to generate molecular markers for L. racemosus to facilitate
the introduction of its useful adaptive genes in wheat breeding.

1.7 Conclusions and Future Prospects

The objectives of wheat breeding programs are linked with the demands of
cumulative population growth and the necessity to increase food production. Due to
the narrow genetic base available in modern bread wheat germplasm, achieving this
objective will be the challenge of the century. Therefore, finding broader sources of
adaptive genes in the wheat gene pool will be essential for introducing traits that
could help increase yields in the warmer, drier climate predicted for most of the
world’s breadbasket. L. racemosus represents a potentially valuable source of
genetic diversity for the improvement of stress tolerance specifically, and wheat
production generally. In this review, we have described L. racemosus genes that
will be potentially useful for improving the tolerance of wheat to biotic and abiotic
stresses and improving its nitrogen use efficiency. However, for the most efficient
exploitation of L. racemosus genes in wheat breeding, the targeted screening of
wheat–L. racemosus cytogenetic stocks for resistance to biotic and abiotic stresses
should be continued. The wealth of molecular genetic information that is becoming
available should be efficiently exploited for breeding purposes, and the shortage
should be replenished through genome wide approaches.

Acknowledgments We thank Dr. M. Kishii, CIMMYT, who gave us valuable information about
L. racemosus addition lines.

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Chapter 2
Arachis Gene Pools and Genetic
Improvement in Groundnut

Anurudh K. Singh and S.N. Nigam

Abstract Groundnut (Arachis hypogaea L.) is an important oilseed and food crop
in the world. The crop is predominantly grown in low input production systems in
developing countries in Asia and Africa. There are several production constraints,
both biotic and abiotic, to groundnut. Some of these are global in nature and the
others are either regional or local. Four Arachis gene pools contain 80 species,
distributed among nine sections are native to five countries of South America.
Section Arachis contains tetraploid cultivated groundnut, divided into two sub-
species and six botanical varieties and a number of cross-compatible diploid species
with rich genetic diversity. International efforts have made significant progress in
collection and conservation of these genetic resources, facilitating genetic
improvement. Groundnut is an autogamous crop. The pedigree and bulk selection
methods are more commonly used by the groundnut breeders. Conventional
breeding, including cytogenetic manipulations introgressing genes from
cross-compatible wild diploid species has been effective in some areas, while in
others it has been tardy due to lack of proper and effective phenotyping tools and
limited understanding of the genomics, genetics/inheritance, and underlying
mechanisms influencing targeted traits. A greater diversification of parental
resources (both cultivated and wild Arachis species) in breeding programs is
required to develop new cultivars with diversified genetic backgrounds, which will
enable them to perform better under the changing climatic/adverse conditions.
Molecular breeding is in infancy. Infrequent and low polymorphisms have
restricted the progress in the development and application of genetic maps, except
in cases where polymorphic chromosomal regions have been introgressed from
diploid wild Arachis species into A. hypogaea. Both conventional and noncon-
ventional crop improvement efforts in groundnut need to concentrate on bridging

A.K. Singh (&)

Plot No. 2924, Sector -23, Gurgaon 122017, Haryana, India
e-mail: [email protected]
S.N. Nigam
Plot No. 125, Road No. 74 Jubilee Hills, Hyderabad 500 033, Telangana, India
e-mail: [email protected]

© Springer International Publishing Switzerland 2016 17

V.R. Rajpal et al. (eds.), Gene Pool Diversity and Crop Improvement,
Sustainable Development and Biodiversity 10,
DOI 10.1007/978-3-319-27096-8_2
18 A.K. Singh and S.N. Nigam

the yield gap between the potential yield and the realized yield by alleviating major
production constraints particularly in rainfed environment.

Keywords Arachis hypogaea Arachis gene pool Center of origin/diversity

Core collection

Genetic improvement

Molecular breeding Genetic

transformation

2.1 Introduction

Groundnut or peanut (Arachis hypogaea L.), an annual legume of indeterminate

growth habit, is primarily grown for its high-quality edible oil (44–56 %) and easily
digestible protein (22–30 %) in its seeds. Groundnut seeds also contain carbohy-
drates (10–25 %) and are a rich source of vitamins (E, K, and B complex), minerals
(Ca, P, Mg, Zn, and Fe), and fiber. It ranks sixth in edible oil production among the
oilseed crops and thirteenth among the food crops (production utilized directly as
food or in confections) in the world. It is also the third most important source of
vegetable protein in the world. It is grown predominantly for food use in North
America, Southern Africa, West Africa, Southeast Asia, and Europe and predom-
inantly for edible oil use in South America and Southwest Asia. In East Africa and
East Asia, both food and edible oil uses are important. On a global basis, 41 % of
the production goes for food use and 49 % for extraction of edible oil. The
remaining 10 % is used as feed and seed or goes waste. The commercial production
of groundnut occurs principally between 40°N and 40°S and about 114 countries
grow groundnut. The crop is predominantly grown in low-input production system
in Asia and Africa with yield ranging between 700 and 1000 kg/ha and in high
input system in USA, Australia, Argentina, Brazil, China, and South Africa where
yields of 2000–4000 kg/ha are obtained.

2.2 Arachis Gene Pool and Taxonomy

Genus Arachis gene pool contains 80 species (Krapovickas and Gregory 1994; Valls
and Simpson 2005). Based on distribution, character clustering, and
cross-compatibilities, Krapovickas and Gregory (1994) classified the genus into nine
sections. Of these, section Trierectoides contains 2, Erectoides 14, Extranervosae 10,
Triseminatae 1, Heteranthae 6, Caulorhizae 2, Procumbentes 10, Rhizomatosae 4,
and Arachis 31 species. Section Arachis contains cultivated groundnut (A. hypogaea),
another tetraploid species A. monticola and 29 diploid wild species. Based on high
degree of genetic isolation and the comparative cytology (Fernandez and Krapovickas
1994), Krapovickas and Gregory (1994) inferred that five sections, Trierectoides,
Erectotoides, Triseminatae, Extranervosae, and Heteranthae, are primitive compared
2 Arachis Gene Pools and Genetic Improvement in Groundnut 19

to sections, Procumbentes, Caulorhizae, Rhizomatosae, and Arachis, except for

section Erectoides. Sections Rhizomatosae and Arachis have evolved feature of
shorter peg length, while sections Caulorrhizae and Rhizomatosae have new methods
of vegetative propagation: stolons and rhizomes, respectively, and appear to be
advanced. Besides, the annual character represents an adaptive advantage that permits
the species to avoid droughts in the northeast Brazil (Heteranthae) and in the foot hills
of the Andes, as well as the flooding of the Paraguay River watershed (Arachis).
Additionally, section Arachis appears to be spreading to new territories invading the
areas of other sections. Its species grow intermixed with populations of Extranervosae
in the upper Paraguay basin and occupy common ground with section Procumbentes
in the Gran Pantanal. They have reached the shores of La Plata and the southeastern
coast of Brazil and grow from Yala in northwest Argentina to the Tocantins in
northeast Brazil, besides worldwide adaptation of A. hypogaea. Cross-compatibility
recorded between A. paraguariensis ssp. paraguariensis, the southernmost taxon in
the section Erectoides, A. rigonii, the westernmost species in the section
Procumbentes, and A. duranensis, one of the most western species of section Arachis
(Gregory and Gregory 1979) and recently between some more species of these sec-
tions (Singh 1998; Mallikarjuna 2005; Mallikarjuna and Hoisington 2009) and A.
glabrata of Rhizomatoseae (Mallikarjuna and Sastri 2002) corroborate the closeness
and advance nature of these sections. Figure 2.1 illustrates the overall phylogenetic
relationships between various sections.

Fig. 2.1 The sections of genus Arachis with connecting lines displaying intersectional
crossability. All species within a section are crossable, except Eurhizomatosae x Prorhizomatosae
20 A.K. Singh and S.N. Nigam

Harlen and de Wet (1971) proposed the gene pool concept in order to provide a
genetic perspective to relationship of cultivated plant species to other components
of genetic diversity, the wild relatives, based on cross-compatibility into (1) primary
gene pool (GP-1), (2) secondary gene pool (GP-2), and tertiary gene pool (GP-3).
Application of this principle facilitates clearer understanding of phylogenetic
relationships between the wild and cultivated species and helps to identify appro-
priate breeding strategies for incorporating desired genes into conventionally usable
form of cultivated species for designing new cultivars. This helps to facilitate
conventional cytogenetic manipulations to establish fertile hybrids, improve genetic
recombination for incorporation of desirable genes into a usable form and
hybridization using pre- or post-fertilization manipulations to establish hybrids.
Alternatively, biotechnological approaches may be applied to access genes through
sexual or parasexual means of genetic transformation. This approach has been used
in groundnut classifying the genetic diversity into four gene pools (Smartt 1990;
Singh and Simpson 1994):
1. The primary gene pool consists of landraces and traditional cultivars of
groundnut from primary and secondary centers of genetic diversity in South
America and other groundnut-growing countries and wild A. monticola found in
northwest Argentina having free crossability with A. hypogaea producing nor-
mal segregants.
2. The secondary gene pool consists of diploid species from section Arachis,
cross-compatible with A. hypogaea, despite ploidy differences, producing sterile
to partially fertile hybrids.
3. The tertiary gene pool includes species of section Procumbentes, which have
crossed with diploid species of section Arachis (Gregory and Gregory 1979;
Mallikarjuna 2005; Mallikarjuna and Hoisington 2009) and probably coevolved
with series perennes of section Arachis; Erectoides, whose species are weakly
cross-compatible with diploid species of section Arachis and A. hypogaea
(Singh 1998); and Rhizomatosae, whose tetraploid species can be crossed both
with diploid species of section Arachis and A. hypogaea (Gregory and Gregory
1979; Mallikarjuna and Sastri 2002).
4. The quaternary gene pool of the remaining Arachis species that are
cross-incompatible or very weakly cross-compatible to species of section
Arachis and are classified into five other sections.
Based on heritable genetic variation observed in cultivated A. hypogaea,
Krapovikas and Gregory (1994) divided it into the following two subspecies and six
botanical varieties.
1. Subsp. hypogaea: Characterized by absence of flowers on main axis and regular
alternation of vegetative and reproductive branches on the laterals and long life
cycle.
2. Subsp. fastigiata: Characterized by presence of flowers on main axis and no
specific order of vegetative and reproductive branches on the laterals and shorter
life cycle.
2 Arachis Gene Pools and Genetic Improvement in Groundnut 21

Subsp. hypogaea is divided into two botanical varieties as follows:

i. Var. hypogaea: Characterized by leaflets with glabrous dorsal surface, short
central or main axis, prostrate to erect growth habit, simple inflorescence, and
2–3 seeded pods.
ii. Var. hirsuta: Characterized by leaflets with entire dorsal surface hairy
(1–2 mm), long central or main axis, prostrate growth habit, and 2–4 seeded
pods.
Subsp. fastigiata is divided into four botanical varieties as follows:
i. Var. fastigiata: Characterized by leaflets with glabrous dorsal surface and hair
only on the midrib, 3–5 seeded pods with smooth or lightly marked reticu-
lation without highlighting the longitudinal ribs, reproductive branches mostly
short and thin, little branched, curved branches, erect growth habit, and simple
inflorescence.
ii. Var. peruviana: Characterized by leaflets with glabrous dorsal surface and hair
only on the midrib, pods with very marked reticulation and with prominent
longitudinal ribs and long and strong reproductive branches (5–10 cm) with
strong central axis and lateral branches.
iii. Var. aequatoriana: Characterized by leaflets with entire dorsal surface hairy
(1–2 mm), long reproductive branches, mainly the lateral branches, central
axis mostly with short inflorescence and reproductive branches, deep pod
reticulation, purple stems, more branched and erect growth habit.
iv. Var. vulgaris: Characterized by pods mostly 2-seeded, bunched fruits pointing
to the base of the plant, erect growth habit, more branched and compound
inflorescence.
Commercially, var. hypogaea is also known as Virginia type (large-seeded) or
Runner type (small-seeded), var. hirsuta as Peruvian runner, var. fastigiata as
Valencia type, and var. vulgaris as Spanish type.
Besides the variability of primary gene pool of cultivated A. hypogaea, the wild
Arachis species have attracted groundnut workers because of their resistance to
diseases and insect pests for which the genetic variation in primary gene pool is
limited. The most accessible variability of primary and secondary gene pools have
been successfully utilized in groundnut improvement and their potential value is
now much more predictable and productive. The exploitation of tertiary and qua-
ternary gene pools waits for advancement in the biotechnological techniques and
policy decision with regard to release of transgenic varieties at global level.
22 A.K. Singh and S.N. Nigam

2.3 Extent of Distribution, Center of Origin, and Genetic

Diversity

The genus Arachis is naturally distributed in east of the Andes, south of Amazon,
north of La Plata from northwest Argentina to northeast Brazil, including
Argentina, Bolivia, Brazil, Paraguay, and Uruguay, i.e., from the mouth of the
Amazon (0°) to south across the Sao Francisco and the Jequitinhonha, and into the
mild temperate zone to 34° S on the shores of the South Atlantic in the southern
Uruguay (Fig. 2.2). Arachis species grow from sea level to 650 m above mean sea
level (amsl) on the Planalto, from southern Mato Grosso to southern Goias, to

Fig. 2.2 Distribution and extent of various sections of genus Arachis in South America (based on
Krapovickas and Gregory 1994/2007)
2 Arachis Gene Pools and Genetic Improvement in Groundnut 23

1450 m near Jujuy. They are found mixed in the vegetation of mixed forest to open
grassland. The species may grow submerged, among stones bathed with water, in
dry gravel and in flood plain alluvium. For these reasons Arachis species are found
from semiarid region to the tropical locations receiving an average rainfall more
than 2000 mm and subjected either to intense drought or flooding. Adaptation of
wild Arachis species to such diverse conditions has resulted in generation of great
genetic variability and resilience to grow under diverse and adverse conditions. This
probably led to the development of geocarpy and tuberiform roots to overcome the
harshness of dry conditions and to escape the seasonal fires. However, in cultivated
groundnut (A. hypogaea), selection pressure against the tubiriform roots led to
elimination of this trait, but the geocarpy providing protection to fruits from adverse
external environment, ensuring regeneration was retained.
Krapovickas and Gregory (1994) considered that the genus Arachis originated in
the Sierra de Amambay, on the border between Mato Grosso do Sul (Brazil) and
Paraguay, where grew, possibly, the oldest species of the genus, A. guaranitica
(Gregory et al. 1980). It has been difficult to understand how the genus could have
dispersed to some 4000 km, both toward northeast up to Amazon as well as to the
west, up to the Andes. Fluvial system associated with rivers, streams and the
deposits, and landforms created by them must have played an important role in
dispersal, as many species have distribution associated with the watershed of the
great Paraguay, Uruguay, and Parana or Sao Francisco rivers. The species generally
live near watercourse, in places where the water evidently reaches only during the
higher floods. The geocarpic habit also indicates possible support to long distance
dispersal of species through the rivers and streams. For this reason, Gregory et al.
(1973, 1980) postulated that most ancient species were found in higher elevations,
their immediate descendants occupied the next lower eroded surfaces, while the
distantly evolved species occupied still lower and more recently eroded surfaces.
Further, as seeds moved to lower elevations, they became isolated in major river
valleys; thus probably different sections of the genus evolved independently in
parallel fashion. This perception, however, is changing with record of overlapped
distribution of species belonging to some sections. Dispersion of species has also
occurred by animal and human movement (Singh et al. 2004).
Based on Krapovickas and Gregory (1994), Fig. 2.2 presents the extent and
distribution of genus Arachis and its various sections in South America.
Section Trierectoides lives in the highest places of the divide between the water-
sheds of the Paraguay and Parana rivers, 400–700 m amsl. The northern limits are
found in Jatai, in Goias at 700 m amsl between the Araguaia and Paranaiba rivers.
Section Erectoides is characteristic of the ‘cerrado’ with red soil, which surrounds
the Mato Grosso Pantanal and is nearly exclusive to Mato Grosso do Sul, with some
species going beyond and others extending into Paraguay. The other group of this
section lives in the southwest extreme of the section’s range. All species of section
Extranervosae live in state of Goias, Tocantins, the central part of Mato Grosso, and
the northern part of the Mining Triangle in Minas Gerais. A few extend beyond these
limits. A. villosulicarpa, the other cultivated species of genus, is grown by the
indigenous people of west central Mato grosso. The majority of the species in this
24 A.K. Singh and S.N. Nigam

section grow on a very special soil type, frequently encountered in the “cerrado”,
constituted by a thin layer of soil over a stony substrate. The lone member of
Triseminatae grows in the state of Bahia, in the south of Pernambuco, in the north of
Minas Gerais, and in the vicinity of the São Francisco River. Section Heteranthae is
a typical of northeastern region of Brazil. Section Caulorrhizae grows in the border
area between the Brazilian states of Goiás, Bahia, and Minas Gerais, reaching as far
as the Atlantic coast, where the type specimen of A. pintoi was collected.
Section Procumentes extends itself along the Paraguay River from Concepción on
the Tropic of Capricorn, toward the north, flanking the Pantanal in Mato Grosso on
the south and the north, and then expands westward as far as Santa Cruz de la Sierra,
in Bolivia, living in soils that are periodically flooded. Section Rhizomatosae’s
tetraploid species occupy a central position within the overall range of the genus
Arachis and the diploid A. burkartii growing more to the south. Section Arachis is
distributed along an axis that coincides more or less with the 57th and 58th
meridians, that encompasses the watersheds of the Paraguay and Uruguay
rivers and ends at the La Plata River. It is the most widely distributed section
invading/overlapping the areas of other sections. The perennial species of section are
found along water house and some are adapted to flooding. Further, two arms of
section extend toward the north and correspond to the basin of the Tocantins River to
the east and to the Mamoré and Guaporé River system to the west, between Trinidad
and Guayaramerín, in Bolivia (Fig. 2.2). In these two expansions, the species
encountered are annuals, adapted to prolonged inundation. A. stenosperma, an
annual growing on the sands of the Atlantic coast, isolated at the eastern extreme,
evidently was carried by the humans (Singh et al. 2004). Expansion of section
toward the southwest is constituted by annual species, adapted to conditions of
periodic drought. They extend from the dry “chaco” up to the first foothills of the
Andes: A. batizocoi (300–950 m amsl) and A. duranensis (250–1250 m amsl)
together with A. monticola (1350–1560 m amsl) grow at the highest elevations.
Highest numbers of species representing eight sections of Arachis are reported
from Brazil, of which four are nearly endemic. Bolivia has the second largest
number of species followed by Paraguay, Argentina, and Uruguay. Most species
occurring in Brazil are confined to the west central region, with a group of species
endemic to the semiarid region of northeast. Further differentiation in patterns of
genetic variability in different sections occurred as a result of their adaptation to
different ecological niche, where they were caught with a series of land uplifts
during their movement downstream in the associated drainage systems. Genetic
isolation among the species of section Arachis is not that strongly marked as among
the species of other sections.
Regarding the origin of cultivated groundnut, A. hypogaea, Krapovickas (1969)
proposed southern Bolivia and northwestern Argentina, which is the range of the
diploid species considered to be involved in its origin. As per Hammons (1994), it
probably first occurred in the valleys of the Parana and Paraguay River systems in
the Gran Chaco area. Krapovickas (1969) suggested the eastern foothills of the
Andes for domestication, based on wide range of ecological diversity and uses of
groundnut. This area is also an important center of diversity of primitive
2 Arachis Gene Pools and Genetic Improvement in Groundnut 25

subsp. hypogaea. Archeological evidence suggests that groundnut has been in

cultivation for over 3500 years. Early European explorers found local Indians
cultivating groundnut in many islands in the Antilles, on the northeast coasts of
Brazil, in all the warm regions of the Rio de la Plata basin, extensively in Peru and
sparsely in Mexico.
Of the two subspecies of A. hypogaea, the primitive subsp. hypogaea has its
most important center of variation in Bolivia. In southeast Bolivia, on the first
foothills of the Andes in the departments of Tarija and Chuquisaca, samples of
cultivated groundnut with the greatest amount of primitive characters have been
collected. A. hypogaea subsp. hypogaea var. hirsuta Köhler was found in the
archeological deposits from the coast of Peru. A. hypogaea subsp. fastigiata var.
fastigiata (Valencia types) has its most important center of variation in Paraguay
and is the most widespread variety in all of South America. A. hypogaea
subsp. fastigiata var. peruviana is grown in almost all of Peru, especially in the
basin of the Marañón River, and is common in Ecuador. Its southern limit is found
in northern Bolivia, where a few samples were found in Rurrenabaque on the Beni
River and in the department of Pando. A few samples were also obtained in Acre
state of Brazil. A. hypogaea subsp. fastigiata var. aequatoriana is nearly confined
to Ecuador, primarily cultivated in the provinces of El Oro and Loja. It is also
cultivated sporadically in northern Peru. A. hypogaea subsp. fastigiata var. vulgaris
(Spanish types) is grown in South America in Uruguay, in Argentina (Santa Fe,
Entre Ríos and Corrientes), in southern Brazil and to some extent, in Paraguay.
Based on above occurrence, Krapovickas (1969) recognized five and Gregory and
Gregory (1976) six centers of genetic diversity, while recent explorations added
Ecuador as the seventh center with distinct group of landraces referred as var.
aequatoriana (Singh and Nigam 1997). These centers (Fig. 2.3) are given below:
1. The eastern foothills of the Andes in Bolivia
2. The Guarani region
3. Goias and Minas Gerais (Brazil)
4. Rondonia and northwest Mato Grosso (Brazil)
5. Peru
6. Northeastern Brazil
7. Ecuador
These centers of diversity present a very high level of genetic variation due to
natural introgressive hybridization between divergent types, followed by human
selection resulting in production of typical hybrid swarms. The first center (eastern
foothills of the Andes in Bolivia) is a center of diversity of subsp. hypogaea var.
hypogaea with few landraces of var. fastigiata. In Bolivia, there are indications of
introgression between the two subspecies, and ‘Overo’ and ‘Cruceno’ types of
groundnut are probably the product of such introgression (Krapovickas 1969;
Gregory and Gregory 1976). The second center of diversity is the Guarani region,
dominated by the erect subsp. fastigiata (Valencia types) with rare occurrence of
subsp. hypogaea and with little evidence of introgression between the two
26 A.K. Singh and S.N. Nigam

Fig. 2.3 Centers of origin and diversity of Arachis hypogaea in South America, I. The eastern
foothills of Andes in Bolivia; II. The Guarani region; III. Goias and Minas Gerais; IV. Rondonia
and northwest Mato Grosso; V. Peru; VI. Northeastern Brazil and VII. Ecuador

subspecies. However, hybrid swarms of intermediate landraces between the two

botanical varieties of subsp. fastigiata, fastigiata and vulgaris, do exist, conse-
quently both var. fastigiata (Valencia), Porto Alegre and var. vulgaris (Spanish)
2 Arachis Gene Pools and Genetic Improvement in Groundnut 27

Negrito are identified. It is possible that spread of Valencia types to other parts of
the world might have occurred from Paruguay or central Brazil, but more likely
point of embarkation is from the northeast coasts. The Guarani region is also center
of diversity for var. vulgaris, the Spanish types, which probably were disseminated
from this region (Krapovickas 1969; Gregory and Gregory 1976). The third center
of genetic diversity, Goias and Minas Gerais, has distinct varietal pattern from the
Guarani, but is still dominated by erect subsp. fastigiata (Valencia types), with very
few representative of subsp. hypogaea. Landraces of botanical varieties, fastigiata
and vulgaris, are found without much indication of introgression. Rondonia, in the
fourth center of diversity represents the nambyquarae types of subsp. hypogaea.
The fifth center of diversity, Peru, is represented by the collection of three
distinct types. One type includes like the one found in pre-Columbian tombs, with
fruits having prominent constriction, veins, and beak and belongs to subsp. hy-
pogaea, referred as Chinese type in USA and another with similar fruit charac-
teristics, but belonging to subsp. fastigiata var. fastigiata. The two together have
been referred as ‘peruvian’ type by Dubard (1906). And a third type with smooth
pods, three to five seeds per pod and almost no beak, belonging to subsp. fastigiata
var. fastigiata has also been collected. The sixth center, the northeast of Brazil, is
regarded as the tertiary center of diversity (Krapovickas 1969; Gregory and Gregory
1976) with almost all botanical types. Seventh center, Ecuador, represents types
similar to var. fastigiata from Peru, but morphologically distinct and might even be
considered as intermediate between vars. fastigiata and hypogaea. Williams (1991)
studied the region of the north Beni of Bolivia/Peru and collected some more
extraordinary types, appearing to be intermediate between subsp. fastigiata and
subsp. hypogaea.
The global dispersal of cultivated groundnut occurred in early 1500, at least in
two distinct forms—a two-seeded Brazilian and three-seeded Peruvian types dis-
persed soon after the discovery of New World (Dubard 1906). Many authorities
believe that the Portuguese carried two-seeded varieties from Brazil to Africa, to the
Malabar Coast of southwestern India and possibly to the Far East. The Peruvian
type (A. hypogaea var. hirsuta) went to the Western Pacific, to China, to Indonesia
(Java), and to Madagascar. Their most plausible path was up the west coast from
Peru to Mexico, thence across the Pacific as an item of trade between Acapulco and
Manila. Gibbons et al. (1972) recorded cultivar clusters of subsp. fastigiata var.
vulgaris representing both the Guariani region and the region of the eastern slopes
of Andes in Bolivia and parts of western Brazil in Africa. In Africa and Asia,
groundnut readapted to environment and specialized agricultural production sys-
tems. Africa received groundnut from Brazil in West Africa and probably from west
coast of South America in the east coast through Philippines, China, and India and
became important center of diversity. The Spanish type of groundnut was intro-
duced into Europe from South America (Krapovickas 1969). Higgins (1951)
speculated that variety hypogaea was introduced to the southeastern United States
from Europe, while Simpson et al. (2001) suggested that cultivated peanut traveled
in slave ships from Africa into the southeastern United States, Central America, and
28 A.K. Singh and S.N. Nigam

northeast South America, thus returning as modified germplasm. By the nineteenth

century, groundnut became an important food crop in West Africa, Southeast and
South Asia, and USA, generating rich genetic diversity.

2.4 Collection, Conservation, and Ensuring Availability

of Genetic Resources

Exploration for collecting seeds and living plants of cultivated groundnut varieties
and wild Arachis species started in mid-twentieth century by USDA and CSIRO
scientists. The first exploration dedicated to collection of germplasm was conducted
in Argentina in 1945 with the initiation of plant breeding program at the Manfredi
Agricultural Experiment Station (Cordoba) and with the organization of the
Department of Plant Exploration and Introduction (DEIP) of the Ministerio de
Agricultura de la Nación, under the direction of E. C. Clos. Since then to early 70s,
extensive explorations were made by Krapovickas (CONICET) and Gregory
(USDA) collecting live specimens of wild species and samples of cultivated
groundnuts. It was followed with introductions of these collections to other parts of
the world. Banks (1976) emphasized the need to make additional collections of the
cultivated groundnut and the wild species before destruction of their habitats.
Consequent to the support of the International Board for Plant Genetic Resources
(IBPGR), FAO, and the International Crops Research Institute for the Semi-Arid
Tropics (ICRISAT), 17 expeditions were undertaken between 1976 and 1983 to the
centers of origin and diversity of Arachis in South America, surveying almost entire
area of distribution of the genus (Valls et al. 1985). These efforts continued,
enriching the available genetic diversity/collections, till the time of enforcement of
Convention on Biological Diversity (CBD), which provided ownership to the
nations of their biological resources and made them responsible for their mainte-
nance and conservation, to facilitate their use. Each expedition provided additional
locations of both wild and cultivated groundnut, as a result the map of natural
resources for different sections was greatly modified from the earlier one presented
by Gregory et al. (1980). Additionally, cultivated groundnut accessions were col-
lected from groundnut-growing areas of various countries, included landraces,
farmers’/traditional varieties, material developed by the breeders and/or released
varieties, and the genetic stocks identified with special features or sources of
resistance to biotic and abiotic stresses, representing different botanical varieties and
cultivar groups. Groundnut germplasm is conserved as pods or seeds, except for
wild Arachis species belonging to section Rhizomatosae, which rarely produce seed
and if produced, progenies are highly heterogeneous and therefore are conserved as
live plants under controlled conditions providing an environment close to their
habitat. Globally, several repositories have facilities for processing and ex situ
conservation/storage of seeds, facilitating prolonged shelf life.
2 Arachis Gene Pools and Genetic Improvement in Groundnut 29

Short-term chambers: These are maintained at 20 ± 5 °C and relative humidity

(RH) 35–45 %. Pods/seeds in these chambers remain viable for several years
without much loss of viability.
Medium-term chambers: These are prefabricated modules maintaining temper-
ature between 4 to 10 °C and RH between 30 to 40 %. The seeds are dried to 8–
10 % moisture level and kept in moisture-proof containers. The seeds remain viable
for 25–35 years without much loss of viability.
Long-term chambers: These are also prefabricated modules maintaining tem-
perature around minus 18 °C and host around 1000–1500 seeds of each accession.
The seeds are dried to a moisture level of 4–5 %, hermetically sealed in vacuum
sealed aluminum pouches, and transferred to the chambers.
The major repositories of the world groundnut germplasm collections are at
ICRISAT, India; in the USA at Southern Regional Plant Introduction in Georgia, at
North Carolina State University, Raleigh, and at Texas Agricultural Experiment
Station, Stephenville; in Brazil at Empressa Brasileria De Pesquisa Agropecuaria
(EMBRAPA)/Centro Nacional de Recursos Geneticos (CENARGEN), Brasilia and
Instituto Agronomico, Campinas; and in Argentina at Instituto Botanica del
Nordeste (IBONE), Corrientes and Instituto Nacional de Technologia Agropecuaria
(INTA), Manfredi. The germplasm collections include both cultivated and wild
Arachis species accessions. ICRISAT maintains over 14,966 from 91 countries,
while USDA and NBPGR (India), 9225 and 13,337 accessions, respectively, and
provide basic genetic stocks to the international and national scientific community.
Table 2.1 presents the germplasm status at some of the major repositories. The low
seed multiplication rate and large seed size of groundnut are the major concerns in
germpasm multiplication and storage.
Despite assembly of large collections, the use of germplasm accessions in
groundnut improvement has remained limited, keeping the genetic base of most
groundnut cultivars narrow and vulnerable to biotic and abiotic stresses and natural
vagaries. Initially, this was circumvented by ensuring the availability and supply of
global genetic resources. For example, ICRISAT since 1980 distributed 38,362
accessions to Indian researchers, leading to increased use of exotic germplasm in
national breeding programs. Consequently, during 1985–1995, 46 % of new
groundnut variety proposals in All India Coordinated Research Project on Oilseeds
had exotic germplasm in their parentage, accruing an increase in national produc-
tion from 794 kg in 1980 to 988 kg in 1994, about 1.4 % per annum (Singh and
Nigam 1996). At the same time to improve information on agronomic and eco-
nomic traits of each accession that requires extensive evaluation and for manage-
able quantification of genetic diversity, core collection concept advocated by Brown
(1989) was applied, leading to creation of a set of 10 % of total collections retaining
most variability of the entire collections. It was hoped that it would facilitate easier
access to genetic resources, enhance their use in crop improvement and also sim-
plify their management in genebank. Initially, core collections were developed by
stratifying germplasm accessions by country of origin and botanical varieties,
followed by the use of data on quantitative morphological traits for principal
component/multivariate analysis and clustering, and randomly selecting 10 % of
30 A.K. Singh and S.N. Nigam

Table 2.1 Groundnut germplasm holding at some important repositories

Repository Number of accessions conserved Additional
Primary gene pool Other gene pools information
ICRISAT, Indiaa Var. hypogaea 6838 + Representing Representing 91
vulgaris Section, Arachis, countries
5493 + fastigiata Rhizomatosae,
2351 + aequitoriana Extranervosae,
14 + peruviana 251 + Erectoides,
hirsuta 19 = 14966 Procumbentes
Caulorhizae,
Triseminatae = 453
GRIN and USAb A. hypogaea Representing Texas A&M,
6804 + ssp. fastigiata Section, Arachis, Experiment Station,
361 + ssp. hypogaea Rhizomatosae, Stephenville,
141 + var. Procumbentes, maintains 1200 acc.
aequatoriana 62 + var. Heteranthae, of 70 wild species,
fastigiata 1149 + var. Extranervosae, 400 hybrids and two
peruviana 24 + var. Erectoides, mapping populations,
vulgaris 128 + var. Trierectoides, while North Carolina
hirsuta 29 + var. Triseminatae = 641 State University,
hypogaea 527 = 9225 USA 740 acc. of
primary and 406 of
other gene pools
NBPGR, Delhi and 13337 at NBPGR, and Representing Represent 90 and 84
DGR, Junagadh, var. hypogaea Arachis, countries
India (personal 2386 + ssp. fastigiata Caulorhizae, respectively, NBPGR
communication) 4458 + others 2280 = Heteranthae, maintains duplicates
9129 at DGR Rhizomatosae, of ICRISAT
Junagadh, India Procumbentes, accession of Indian
Erectoides = 112, origin
105 respectively
Oil Crops Research 7837 + 4210 246 Core of 576 and
Institute, Chinese mini-core of 298
Academy of
Agricultural
Sciences, and Crops
Research Institute,
Guaeglong Academy
of Agricultural
Sciences, Chinac
CENARGEN, Brazil 1200 + 2140 Representing
and Instituto Section, Arachis,
Agronomicas Caulorhizae,
Campinas, Brazilc Rhizomatosae,
Extranervosae,
Erectoides,
Triseminatae
= 1220
INTA, Manfredi and 3534 106 + 472
IBONE, Argentinac
Source
a
//ICRISAT// Groundnut Crop: www.icrisat.org/crop-groundnut-genebank.htm
b
www.ars-grin.gov/npgs/cgc_reports/Status11.pdf
c
Pandey et al. (2012)
2 Arachis Gene Pools and Genetic Improvement in Groundnut 31

collections to constitute a core collection. Using this methodology, first core col-
lection of 831 accessions was developed on US germplasm collection of 7,432 by
Holbrook et al. (1993); followed by the development of a core collection of 1,704
accessions on 14,310 world collections assembled at ICRISAT (Upadhyaya et al.
2003); a core collection of 576 from a collection of 6,390 accessions in China
(Jiang et al. 2007) and another core collection of 576 accessions, and a mini-core
collection of 298 accessions from a collection of 6,839 conserved at the Oil Crops
Research Institute of Chinese Academy of Agricultural Sciences at Wuhan (Jiang
et al. 2013). Using statistical methods it was ensured that these core collections
retained the diversity index and phenotypic correlation of different traits to that of
the entire collections so that they represented most spectrum of variability and were
effective in the genetic improvement of groundnut. These core collections were
further evaluated in multilocations for identification of regional core (Upadhyaya
et al. 2005), development of mini-cores (Upadhyaya et al. 2002a), for identification
of variability for specific traits (Upadhyaya et al. 2006; Jiang et al. 2013); char-
acterizing the core collections using specific molecular markers to enable better
quantification of genetic variability (Kottapalli et al. 2007), and identifying
accessions with specific trait/resistance associated with molecular markers
(Chamberlin et al. 2010). These efforts have been extended to characterization of
diversity using association mapping for exploring the molecular basis of phenotypic
variations, demonstrating a great potential of integrating the association analysis
and marker-assisted breeding by utilizing the mini-core collection (Ren et al. 2014).
Attempts are also made to purify the accessions of mini-core and register them on
the basis of morphological, biochemical, and resistance traits (Chen et al. 2013a).
Comparison of core collections developed in different parts of the world showed
different traits contributing to variability in different set of collections, associated to
the dominance of subspecies and botanical varieties in a collection and selection
pressure (Jiang et al. 2008), indicating want of a universal core collection for
groundnut improvement meeting everyone needs. Most cores are proportionally
limited in variability from vars. hirsuta, peruviana, and aequatoriana.

2.5 Major Constraints to Groundnut Production

Groundnut suffers from several biotic and abiotic production constraints. Some of
them are global in nature; and others are either regional or local.
Biotic constraints: Among the foliar fungal diseases, early leaf spot (ELS;
Cercospora arachidicola Hori.), late leaf spot [LLS; Phaeoisariopsis personata
(Berk.&M.A. Curtis) Van Arx], and rust (Puccinia arachidis Spegazzini) are wide
spread and are prevalent wherever groundnut is grown. Other foliar fungal diseases,
which could be important in certain regions/countries, include web blotch (Phoma
arachidicola Marasas, Pauer & Boerema) and pepper spot and leaf scorch
[Leptosphaerulina crassiasca (sechet) Jackson &Bell]. Among the seed and
seedling fungal diseases, preemergence seed and seedling rots [Aspergillus flavus
32 A.K. Singh and S.N. Nigam

Link ex Fries, A. niger van Tieghem, Macrophomina phaseolina (Tassi) Goidanich,

Sclerotium rolfsii Saccardo, Rhizoctonia solani Kühn, Lasiodiplodia theobromae
(Pat.) Griffon & Maubl., Rhizopus spp., Penicillium spp., Phythium spp., and
Fusarium spp.], Aspergillus crown rot/ collar rot (A. niger van Tieghem), yellow
mold (A. flavus Link ex Fries), and Rhizoctonia damping off (Rhizoctonia solani
Kühn) are wide spread. Important stem, root, and pod diseases caused by fungi
include stem and pod rots (Sclerotium rolfsii Saccardo), Sclerotinia blight
[Sclerotinia sclerotiorum (Lib.) de Bary], Cylindrocladium black rot [CBR;
Cylindrocladium crotalariae (Loos) Bell & Sobers], Botrytis blight (Botrytis
cinerea Pers. Ex Fries), Fusarium wilt (Fusarium oxysporum Schlechtend. Emend
Snyder & Hans.), charcoal rot [Macrophomina phaseolina (Tassi) Goidanich], and
pod rot [Pythium myriotylum Dreschler, Rhizoctonia solani Kühn, Fusarium solani
(Mart.) Saccardo f. sp phaseoli (Burkholder) Snyder & Hans., Fusarium oxysporum
Schlechtend. Emend Snyder & Hans., and Macrophomina phaseolina (Tassi)
Goidanich] are important. The groundnut pod and kernels can also get infected
while developing with A. flavus Link ex Fries/A. parasiticus Speare leading to their
contamination by aflatoxin. The only bacterial disease of significance is bacterial
wilt [BW; Ralstonia solanacearum (E.F. Smith)], which is wide spread in East and
Southeast Asia.
Significant virus diseases include peanut stripe (PStV; Peanut stripe virus) in
East and Southeast Asia, peanut clump (PCV; Peanut clump virus) in West Africa,
peanut bud necrosis (PBND; Peanut bud necrosis virus) in South Asia, tomato
spotted wilt (TSWV; Tomato spotted wilt virus) in North America, peanut stem
necrosis (PSND; Tobacco streak virus) in South India, and groundnut rosette dis-
ease (GRD; a complex of groundnut rosette virus, groundnut rosette assistor virus,
and a satellite RNA) in Africa. Important diseases caused by nematodes are root
knot [Meloidogyne arenaria (Neal) Chitwood, M. hapla Chitwood, M. javanica,
and M. incognita; the first two are wide spread], root lesion [Pratylenchus
brachyurus (Godfrey) Filipjev & Sch. Stekh.], and Kalahasti malady
(Tylenchorhynchus brevilineatus Williams) in Andhra Pradesh in India.
Defoliators, tobacco caterpillar/tobacco armyworm (Spodoptera litura Fab.), hairy
caterpillars (Amsacta albistriga Walk., A. moori Butler), Bihar hairy caterpillar
[Spilosoma (Diacrisia) oblique (Walk.)], gram pod borer (Helicoverpa armigera
Hübner) and groundnut leaf miner (Aproaerema modicella Deventer), sucking pests,
aphids (Aphis craccivora Koch.), thrips (Scirtothrips dorsalis Hood., Thrips palmi
Karny., Frankliniella schultzei Trybom) and Caliothrips indicus Bagnall and
jassids/leaf hopper (Empoasca kerri Pruthi), root and pod feeders, white grub
[Lachnosterna (= Holotrichia) consanguinea Blanch.] and L. serrata (Fab.),
termites/white ants (Microtermes spp. and Odontotermes spp.) and earwig (Anisolabis
stali Dohrn) and storage pests groundnut, bruchid (Caryedon serratus Oliver), and red
flour beetle (Tribolium castaneum Herbst) are the major insect pests of groundnut.
Abiotic constraints: As a majority of groundnut is grown under rainfed condi-
tions, drought is the most significant abiotic stress affecting groundnut production.
Drought can occur at any stage—early-season, mid-season, end-of-season, and
intermittent. Drought also predisposes groundnut pods to aflatoxin contamination
2 Arachis Gene Pools and Genetic Improvement in Groundnut 33

by A. flavus. Other abiotic constraints include low soil fertility, salinity, iron
chlorosis, aluminum toxicity, cold temperature at germination, and high tempera-
ture at podding and harvest.

2.6 Searching Potential Genetic Resources and Managing

Constraints with Genetic Improvement

For assessment of genetic variability and identification of genetic resources with

desired features, the groundnut germplasm assembled at various places have been
characterized and evaluated based on the common groundnut descriptors developed
by IBPGR/ICRISAT (1992). Variability analysis has shown greater variation in
landraces and in the accessions collected from the primary and secondary centers of
diversity in South America, particularly for resistance to biotic and abiotic stresses
and agronomic features like seed mass (Singh and Simpson 1994). An assessment
of genetic diversity on world collection at ICRISAT for 16 morphological and 10
agronomic traits has shown vast diversity in size and shape of pods and seeds.
Principal Component analysis using 38 traits and clustering on first seven PC scores
produced three clusters; consisting North America, middle East, and East Asia in
the first cluster, South America in the second cluster, and West Africa, Europe,
Central Africa, South Asia, Oceania, Southern Africa, Eastern Africa, Southeast
Asia, Central America, and Caribbean in the third cluster. The means for different
agronomic traits differed significantly among regions, while the variances for all the
traits among regions were heterogeneous. South America cluster showed 100
percent range variation for 12 of the 16 morphological traits and revealed highest
range of variation. Assessment of phenotypic diversity in core collection revealed
significant variation. The average phenotypic diversity index was higher in the
fastigiata group (0.146) than the hypogaea group (0.141). The hypogaea group
showed significantly greater mean pod length, pod width, seed length, seed width,
yield per plant, and 100-seed weight than the fastigiata group in both rainy and
postrainy seasons whereas it was opposite for plant height, leaflet length, leaflet
width, and shelling percentage with fastigiata group showing significantly greater
means. Principal coordinate and principal component analyses showed that 12
morphological descriptors and 15 agronomic traits were important in explaining
multivariate polymorphism. Leaflet shape and surface, color of standard petal
markings, seed color pattern, seed width, and protein content did not significantly
account for variation in the first five principal coordinates or components of fasti-
giata and hypogaea types, indicating their relatively low importance. The average
phenotypic diversity index was similar in both subspecies. The Shannon–Weaver
diversity index varied among traits between the two and the diversity within a
subspecies/group depended upon the season and traits recorded. Molecular profiling
of joint composite collection developed by ICRISAT and EMBRAPA, using 21
SSRs showed rich allelic diversity, group-specific unique alleles, and common
34 A.K. Singh and S.N. Nigam

alleles sharing between subspecies and geographical groups. Gene diversity ranged
from 0.559–0.926, with an average of 0.819. Group-specific unique alleles were
101 in wild Arachis, 50 in subsp. fastigiata, and only 11 in subsp. hypogaea.
Accessions from Americas revealed the highest number of unique alleles (109),
while Africa and Asia had only six and nine, respectively. The two subsp. hypogaea
and fastigiata shared 70 alleles. In contrast, the wild Arachis shared only 15 alleles
with hypogaea and 32 alleles with fastigiata (//ICRISAT// Groundnut Crop: www.
icrisat.org/crop-groundnut-genebank.htm). Greater genetic diversity among the
landraces originating from primary and secondary centers of diversity in South
America is corroborated by the molecular characterization using various markers,
for example, in Bolivian landraces (Husain and Mallikarjuna 2012).
For many traits, the primary gene pool has been found limited, but the wild
Arachis species have been found with desired variability; for example, for PStV no
resistant line was found in cultivated groundnut despite screening of 9,000 acces-
sions, but several accessions of wild Arachis showed negative reaction (Culver
et al. 1987; Prasada Rao et al. 1991). Often wild Arachis species have shown higher
level of resistance than primary gene pool. Variability observed among the
accessions of wild species for their reaction against specific constraints (Singh et al.
1996) demands thorough investigation for useful exploitation. Table 2.2 presents
the number of accessions identified with useful diversity and used in breeding
programs, and Table 2.3 lists representative wild species with multiple resistances.

2.6.1 Genetic Improvement Using Resources of Primary

Gene Pool Through Conventional Breeding

Groundnut is a highly self-pollinated crop, though cross-pollination can reach as

high as 10 % at locations and in seasons, where bee activity is high. Standard
breeding methods, followed in groundnut for developing a cultivar, can be placed
into two groups—(a) methods without hybridization, and (b) methods after
hybridization. The former includes introduction, pure line/ mass selection and
mutation breeding and the latter bulk selection, pedigree selection, bulk pedigree
selection, single seed descent method, backcross method, and recurrent selection.
Among these, pedigree and bulk selection methods are more commonly used by the
breeders. Some breeders use a combination of breeding methods or make modifi-
cation to conventional methods.
Examples of release of cultivars following introduction and selection include,
among others, release of JL 24 in India, release of Indian cultivars, TMV 2 and JL
24, under different names in many countries in Southeast Asia and Africa, release of
Makulu Red, Apollo, Egret, Chalimbana, Mani Pintar, and Malimba in Africa, and
New Mexico Valencia C in the USA. Mutation breeding uses X-rays, gamma rays,
and various chemicals to create mutations breaking specific linkages and enhancing
variation for specific character in a genotype. Bhabha Atomic Research Centre
2 Arachis Gene Pools and Genetic Improvement in Groundnut 35

Table 2.2 Primary Gene Pool Genetic diversity for useful traits in world collections at ICRISAT
Stress/trait Acc. Acc. with desirable Additional information
screened variability
Fungal and bacterial diseases
Early leaf spot 7000 37 (2) 15 (India) + 4 (Malawi) + 18
(West Africa)
Late leaf spot 13000 69 (26)
Rust 13000 169 (35)
Aflatoxin 582 4
production
Aflatoxin seed 580 39 (4)
invasion
Pod and stem rot 3222 24/9 (6) 4 (India) + 5 (USA)
Bacterial wilt 24 Screening in Indonesia and
China
Viral diseases
PBNV 7400 23
PMV 6944 2 Promise in wild Arachis spp.
Insect pest
Thrips 5345 15 (7) Promise in wild Arachis spp.
Jassids 136 30/6 (7)
Termites 520 20/9 (6)
Leaf minor 600 18/4 (6)
Aphids 300 4 (1)
Abiotic stress
Drought 820 46 (8)
N fixation 342 4 (2)
Nutritional traits
High oil content 8868 20/44 (10)
High protein 8868 117/51
content
Source Singh and Nigam (1997), Singh et al. (1997) Figure in parenthesis indicate number of
commonly used source in breeding program

(BARC), Mumbai, India used gamma rays to create desired variation for further use
in conjunction with other breeding methods and released 15 groundnut cultivars till
date. Some of these are TG #s 19, 37A, 38B and 51, TAG 24, TGB 39, TPG 41,
TLG 45.
Hybridization provides opportunity to combine genes from different parents and
recombine them in a single genotype via single cross, three-way cross, four-way
cross, convergent cross, diallel mating, and diallel selective mating. Selection is
practiced in segregating generations following the method of selection of choice.
Generally, this process takes 12–15 years, but can be expedited by taking multiple
crops in a year under controlled greenhouse conditions or raising off-season nursery
at other locations where environmental conditions are favorable to raise a crop.
36 A.K. Singh and S.N. Nigam

Table 2.3 Representative desirable genetic diversity in secondary, tertiary, and quaternary gene
pool of Arachis
Species/gene poola Early Late Rust PStVb GRDc Thrips Leaf Lepidoptera
leafspot leafspot hopper
Secondary gene pool (Sect. Arachis)
A. diogoi R R R
A. duranensis MR – I I – S I HR
A. spegazzini MR _ I R – R HR HR
A stenosperma HR HR HR – R HR HR HR
A. villosa R R I S R – – –
A. correntina – – I R – HR HR HR
A. cardenasii HR HR I R R HR HR HR
A. chacoense HR HR I S – HR HR HR
A. kempff- R R – – – – R R
mercadoi
Tertiary gene pool
A. appressipila (P) R – R – R – – –
A. rigonii (P) – – – S – HR I HR
A. benthamii (E) MR MR – I or R – – – –
A. paraguariensis R MR R R – – R –
(E)
A. glabrata (R2) S or MR S or I R or I – I HR HR
MR
Quaternary gene pool
A. repens (C) R R R I – I HR HR
A. lutescens (Ex) R HR –
B. macedoi (Ex) R – – – – I I –
A. villosulicarpa HR HR I – – – – –
(Ex)
A. pusilla (Tri) R R I – – I I HR
A. triseminata R R R R R R
(Tri)
Source Stalker and Moss (1987), Upadhyaya et al. (2011)
a
Symbol in parenthesis for section; Reaction- MR Moderately resistant, R Resistance, HR Highly
resistant, I Immune, S Susceptible, it may vary between accessions of same species
b
PStV = Peanut stunt virus
c
GRD = Groundnut rosette disease

In spite of limited DNA polymorphism, there is abundant morphological vari-

ability present for most of the traits among germplasm accessions of the cultivated
species. However, only an insignificant portion of this large variability has been
utilized for crop improvement for reasons described earlier. To promote intensified
and diversified use of genetic resources in crop improvement, recently, core and
mini-core collections in groundnut have been developed (Holbrook et al. 1993;
2 Arachis Gene Pools and Genetic Improvement in Groundnut 37

Upadhyaya et al. 2002a, 2003, 2010), which capture representative variability in

cultivated groundnut germplasm collection. However, in any applied breeding
program where breeders have to maintain physical, chemical, and esthetic quality
parameters as per the market requirements, breeders are often reluctant to use
primitive germplasm because of linkage drag, which takes a long time to get rid off.
Any variation in these qualities discourages processors to accept new genotypes as
these variations affect the quality of their products and efficiency of their processing
operations. Having used primitive germplasm in the beginning of the program, the
breeders prefer to use second- or third-generation breeding lines with desired genes
for use in breeding programs. In a recent publication, Janila and Nigam (2013) have
reviewed the phenotyping protocols for various biotic and abiotic stresses, which
are being followed in groundnut improvement programs. Murthy and Reddy (1993)
and Reddy and Murthy (1996) have summarized the results of various genetic and
inheritance studies covering most of the traits in groundnut. For details, readers are
advised to refer to these publications. The status of up-to-date efforts, made for
genetic improvement of groundnut in relation to various traits of significance for an
overall engineering of cultivars incorporating both desirable agronomic features and
resilience to stress factors, is described below.
Yield and yield-related traits: Yield is a complex trait with quantitative inheri-
tance. In addition to yield, pod and seed characters are also important for esthetic
and commercial considerations. Most of the pod and seed characters with few
exceptions are governed by a few genes. Selection either for higher pod yield or for
greater harvest index is essential for improvement of yield potential in future cul-
tivars. Remobilization of reserves from vegetative biomass to pods under conditions
of source limitations (falling temperature, defoliation by pathogens or water stress)
is likely to be significant in maintaining yields but may limit response to improved
conditions specially with high partitioning. Newer high-yielding cultivars in the
USA allocate a greater proportion of biomass to reproductive tissue early in the
growth cycle with greater reproductive efficiency and have more spreading growth
habit and greater seed and pod weight than older cultivars (Wells et al. 1991; Seaton
et al. 1992). However, the crop duration in the USA is much longer (140–160 days)
than the one available to the crop in South and Southeast Asia and West Africa
(<100 days).
Between 1944 and 1987, the average yearly genetic gain for yield in Virginia
market type cultivars in the USA was 14.7 kg per ha. However, when the emphasis
in breeding programs shifted to pest resistance, earliness, and quality, the new
cultivars improved upon these traits but failed to combine them with increased yield
potential (Mozingo et al. 1987). During 1980s and mid-1990s, the groundnut yield
in India increased by 1.4 % per annum (Nigam et al. 1991). New Spanish-type
cultivars in India have greater seed size, seed weight, and pod numbers per plant
than the older cultivars (Rathnakumar et al. 2012). Increase of 0.43 % per year in
seed yield, of 0.29 % per year in seed weight, and of 0.52 % per year in pod growth
rate during 1948–2004 were obtained in Argentina (Haro et al. 2013).
Resistance to foliar fungal diseases: ELS, LLS, and rust are the most widely
distributed and economically important diseases of groundnut in the world. Only
38 A.K. Singh and S.N. Nigam

one leaf spot dominates in a region, however, both pathogens can be observed in
the same field. LLS and rust often occur together. Breeding for resistance to foliar
diseases in groundnut got a real boost in the late 1970s and early 1980s when a
massive field screening program for resistance to foliar fungal diseases (rust and
LLS) of more than 13,000 germplasm accessions from 89 countries was launched at
ICRISAT, Patancheru, India. Subsequently, many sources of resistance were
identified and used in breeding programs (Singh et al. 1997). Most of these resistant
sources are landraces from South America and have undesirable agronomic char-
acters (low yield, poor pod and seed traits, and longer crop duration). The com-
ponents of resistance include longer incubation period and latent period, reduced
sporulation, smaller lesion diameter, lower infection frequency, and less defoliation
in resistance sources. Combining high levels of resistance to ELS and LLS into
high-yielding cultivars with acceptable market traits continues to be difficult.
The first-generation cultivars emanating from foliar diseases resistance programs
in India, ICG (FDRS) 10 and Girnar 1, did not find acceptance among the farmers
and traders in spite of their higher yields under heavy disease pressure due to
unattractive pod and seed characteristics. However, when these cultivars were
recycled again in breeding programs, the resultant second-generation genotypes had
better pod and seed characteristics and more acceptability among farmers and
traders in spite of some dilution in their levels of resistance.
ELS, is more serious in Southern Africa and the USA. Resistant/tolerant A.
hypogaea genotypes have been identified in Malawi, West Africa, India, and the
USA with a disease score ranging between 3.6 and 6.3 on a 1–9 scale, where 1 = no
disease and 9 = more than 81 % foliage destroyed. However, resistant sources
reported from the USA (NC 3033 and PI #s 270806, 259747, and 350680) were
found susceptible in India and Malawi. Excessive use of chemicals to control the
disease in the USA is suspected to have led to variation in pathogen.
Genotypes ICG #s 6284, 6902, 7878, 10000, 10948, and 13917 show some
resistance at more than one location. Rate-reducing resistance is quantitative in
nature and controlled by both additive and nonadditive gene effects including
maternal effects. Duplicate recessive inheritance is also observed. Narrow sense
heritability varies from low to high. Some of the ELS tolerant cultivars released in
India are ICGS 44, ICGS 76, M 335, BG 3, Somnath, CSMG 84-1, M 522, Prutha,
and GG 7 and in the USA are VA 81B, DP 1, Georganic, C-99R, Bailey, Florida
07, and Sugg.
LLS is predominant in warmer areas. Sixty-nine A. hypogaea genotypes tolerant
to LLS with disease score ranging between 3 and 5 on a 1–9 scale (described
earlier) have been identified. Forty-nine of these resistant sources are landraces
belonging to var. peruviana with low yield and shelling outturn and other unde-
sirable pod and seed characters. Resistance to LLS is quantitative in nature and
governed by both additive and nonadditive gene effects including maternal effects.
Duplicate recessive inheritance is also reported. Tolerant cultivars released in India
include RG 141, ICG(FDRS) 10, ICGV #s 86590 and 86325, K 134, Girnar 1,
GBPD 4, R 8808, ALR #s 1, 2, and 3, BSR 1, VRI 5, and CSMG 84-1 and in the
2 Arachis Gene Pools and Genetic Improvement in Groundnut 39

USA, Southern Runner, Florida MDR 98, TUFRunner TM ‘727’, Florida 07, and
C-99R, among others.
For rust, of the 169 A. hypogaea genotypes reported resistant (a score of five or
less on a 1–9 scale), 135 are landraces belonging to var. peruviana. Many of these
(ICG #s 7896, 7897, 7899, 10014, 10030, 10052, 10053, 10067, 10933, 10939,
10940, and 10943) have a disease score of <3 but are agronomically poor (low
shelling outturn, thick pod shell, strong pod reticulation, and unacceptable seed coat
color). New sources of resistance—ICG #s 10056, 10567, 10925, 10932, 11108,
12059, 12112, and 12113 and the interspecific derivatives involving A. batizocoi
and A. duranensis—have high levels of resistance with good agronomic potential
and resistance to other biotic stresses. Resistance to rust is reported to be recessive,
partial dominant, or dominant with duplicate recessive, digenic, trigenic, or
multigenic inheritance. Resistant cultivars released in India include ICG (FDRS)
10, ICGV 86590, and GBPD 4, among others.
Resistance to rust and LLS is correlated (r = 0.48–0.60). Forty-two LLS resistant
genotypes are also resistant to rust. Of these, ICG #s 1703, 4995, and 10920 and
interspecific derivative ICG 13917 [259-2 (red)] are useful in multiple resistance
breeding, the last one being resistant to all the three pathogens. Other useful sources
of resistance to both LLS and rust with agronomic potential are A. hypogaea
genotypes ICG #s 6330, 7884, 10023, 10035, and 11182 and interspecific
derivatives ICG #s 11312, 11317 (also resistant to ELS), 11321, 11325, 11337,
13916, 13917, 13919, 13920, and 13922. Cultivars reported with multiple resis-
tances to foliar diseases, among others, are ICG (FDRS) 4, ICG (FDRS) 10, Girnar
1, ICGV 86590, Somnath, GBPD 4, VRI 2, VRIGn 5, and ALR #s 1, 2, and 3 in
India and Azivivi, Nkosour, Adepa, and Jenkaa in Ghana.
Resistance to soil-borne fungal diseases: Breeding for resistance to soil-borne
fungal diseases continues to be difficult as creating uniform disease pressure in the
disease screening nursery remains challenging. Breeding for resistance to A.
flavus/A. parasiticus and aflatoxin contamination has received the most attention
among the soil-borne fungal diseases. Other diseases, where breeding efforts are in
progress, include pod and stem rots, cylindrocladium black rot (CBR), and scle-
rotinia blight.
Efforts on breeding for resistance to A. flavus/A. parasiticus invasion and afla-
toxin production in the USA, where A. parasiticus is dominant, and in other
countries in Asia and Africa, where A. flavus predominates, are in progress.
However, they have not yet succeeded in ensuring complete freedom from A.
flavus/A. parasiticus infection and aflatoxin contamination in groundnut cultivars.
Liang et al. (2009) and Nigam et al. (2009) have reviewed the progress in breeding
for resistance to A. flavus invasion and aflatoxin contamination at ICRISAT,
India/West Africa and Guangdong Academy of Agricultural Sciences in China,
respectively.
There are three barriers to A. flavus/A. parasiticus infection and aflatoxin pro-
duction in groundnut seed—pod wall, seed coat, and cotyledons. Resistance to pod
infection is attributed to shell wall structure and that of seed coat to thickness and
density of palisade layers, absence of fissures and cavities, and presence of wax and
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could be thought of to take Townshend’s place, and Lord North, who
never liked to offend the king, declined the appointment. Before
Chatham could devise a way out of his quandary, his malady again
laid him prostrate, and Townshend was not only not turned out, but
was left practically supreme in the cabinet. The new measures for
taxing America were soon passed. In the debates on the Stamp Act,
it had been argued that while Parliament had no right to impose a
direct tax upon the Americans, it might still properly regulate
American trade by port duties. The distinction had been insisted upon
by Pitt, and had been virtually acknowledged by the Americans; who
had from time to time submitted to acts of Parliament imposing
duties upon merchandise imported into the colonies. The
Nay, more, when charged with inconsistency for Townshend
submitting to such acts while resisting the Stamp Act, Acts
several leading Americans had explicitly adopted the
distinction between internal and external taxation, and declared
themselves ready to submit to the latter while determined to resist
the former. Townshend was now ready, as he declared, to take them
at their word. By way of doing so, he began by laughing to scorn the
distinction between internal and external taxation, and declaring that
Parliament possessed the undoubted right of taxing the Americans
without their own consent; but since objections had been raised to a
direct tax, he was willing to resort to port duties,—a measure to
which the Americans were logically bound to assent. Duties were
accordingly imposed on wine, oil, and fruits, if carried directly to
America from Spain or Portugal; on glass, paper, lead, and painters’
colours; and lastly on tea. The revenue to be derived from these
duties was to be devoted to paying a fixed salary to the royal
governors and to the justices appointed at the king’s pleasure. The
Crown was also empowered to create a general civil list in every
colony, and to grant salaries and pensions at its arbitrary will. A board
of revenue commissioners for the whole country was to be
established at Boston, armed with extraordinary powers; and general
writs of assistance were expressly legalized and permitted.
 HOUSE OF COMMONS
Such was the way in which Townshend proceeded to take the
Americans at their word. His course was a distinct warning to the
Americans that, if they yielded now, they might expect some new
Stamp Act or other measures of direct taxation to follow; and so it
simply invited resistance. That no doubt might be left on this point,
the purpose for which the revenue was to be used showed clearly
that the object of the legislation was not to regulate trade, but to
assert British supremacy over the colonies at the expense of their
political freedom. By providing for a civil list in each colony, to be
responsible only to the Crown, it aimed at American self-government
even a more deadly blow than had been aimed at it by the Stamp
Act. It meddled with the “internal police” of every
Attack on
colony, and would thus have introduced a most the New
vexatious form of tyranny as soon as it had taken York
effect. A special act by which the Townshend revenue assembly
acts were accompanied still further revealed the
temper and purposes of the British government. The colony of New
York had been required to provide certain supplies for the regular
troops quartered in the city, under command of General Gage; and
the colonial assembly had insisted upon providing these supplies in its
own way, and in disregard of special instructions from England. For
this offence, Parliament now passed an act suspending the New York
assembly from its legislative functions until it should have complied
with the instructions regarding the supplies to the army. It need not
be said that the precedent involved in this act, if once admitted,
would have virtually annulled the legislative independence of every
one of the colonial assemblies.

HOUSE OF LORDS
We may perhaps wonder that a British Parliament
Parliament
should have been prevailed on to pass such audacious did not
acts as these, and by large majorities. But we must properly
remember that in those days the English system of represent
representation was so imperfect, and had come to be the British
people
so overgrown with abuses, that an act of Parliament
was by no means sure to represent the average judgment of the
people. The House of Commons was so far under the corrupt
influence of the aristocracy, and was so inadequately controlled by
popular opinion, that at almost any time it was possible for an
eloquent, determined, and unscrupulous minister to carry measures
through it such as could never have been carried through any of the
reformed Parliaments since 1832. It is not easy, perhaps, to say with
confidence what the popular feeling in England was in 1767 with
reference to the policy of Charles Townshend. The rural population
was much more ignorant than it is to-day, and its political opinions
were strongly influenced by the country squires,—a worthy set of
men, but not generally distinguished for the flexibility of their minds
or the breadth of their views. But as a sample of the most intelligent
popular feeling in England at that time, it will probably not be unfair
to cite that of the city of London, which was usually found arrayed on
the side of free government. No wiser advice was heard in
Parliament, on the subject of the New York dispute, than was given
by Alderman Beckford, father of the illustrious author of Vathek,
when he said, “Do like the best of physicians, and heal the disease by
doing nothing.” On many other important occasions in the course of
this unfortunate quarrel, the city of London gave expression to
opinions which the king and Parliament would have done well to
heed. But even if the House of Commons had reflected popular
feeling in 1767 as clearly as it has done since 1832, it is by no means
sure that it would have known how to deal successfully with the
American question. The problem was really a new one in political
history; and there was no adequate precedent to guide the
statesmen in dealing with the peculiar combination of considerations
it involved. As far as concerned the relations of Englishmen in
England to the Crown and to Parliament, the British Constitution had
at last reached a point where it worked quite smoothly. All
contingencies likely to arise seemed to have been provided for. But
when it came to the relations of Englishmen in America to the Crown
and to Parliament, the case was very different. The case had its
peculiar conditions, which the British Constitution in skilful hands
would no doubt have proved elastic enough to satisfy; but just at this
time the British Constitution happened to be in very unskilful hands,
and wholly failed to meet the exigencies of the occasion. The chief
difficulty lay in the fact that while on the one hand the
Difficulty of
American principle of no taxation without the problem
representation was unquestionably sound and just, on
the other hand the exemption of any part of the British Empire from
the jurisdiction of Parliament seemed equivalent to destroying the
political unity of the empire. This could not but seem to any English
statesman a most lamentable result, and no English statesman felt
this more strongly than Lord Chatham.
There were only two possible ways in which the difference could
be accommodated. Either the American colonies must elect
representatives to the Parliament at Westminster; or else the right of
levying taxes must be left where it already resided, in their own
legislative bodies. The first alternative was seriously considered by
eminent political thinkers, both in England and America. In England it
was favourably regarded by Adam Smith, and in America by Benjamin
Franklin and James Otis. In 1774, some of the loyalists in the first
Continental Congress recommended such a scheme. In
Representati
1778, after the overthrow of Burgoyne, the king on of
himself began to think favourably of such a way out of Americans
the quarrel. But this alternative was doubtless from the in
first quite visionary and unpractical. The difficulties in Parliament
the way of securing anything like equality of
representation would probably have been insuperable; and the
difficulty in dividing jurisdiction fairly between the local colonial
legislature and the American contingent in the Parliament at
Westminster would far have exceeded any of the difficulties that have
arisen in the attempt to adjust the relations of the several States to
the general government in our Federal Union. Mere distance, too,
which even to-day would go far toward rendering such a scheme
impracticable, would have been a still more fatal obstacle in the days
of Chatham and Townshend. If, even with the vast enlargement of
the political horizon which our hundred years’ experience of
federalism has effected, the difficulty of such a union still seems so
great, we may be sure it would have proved quite insuperable then.
The only practicable solution would have been the frank and cordial
admission, by the British government, of the essential soundness of
the American position, that, in accordance with the entire spirit of the
English Constitution, the right of levying taxes in America resided
only in the colonial legislatures, in which alone could American
freemen be adequately represented. Nor was there really any reason
to fear that such a step would imperil the unity of the empire. How
mistaken this fear was, on the part of English statesmen, is best
shown by the fact that, in her liberal and enlightened dealings with
her colonies at the present day, England has consistently adopted the
very course of action which alone would have conciliated such men
as Samuel Adams in the days of the Stamp Act. By pursuing such a
policy, the British government has to-day a genuine hold upon the
affections of its pioneers in Australia and New Zealand and Africa. If
such a statesman as Gladstone could have dealt freely with the
American question during the twelve years following the Peace of
Paris, the history of that time need not have been the pitiable story of
a blind and obstinate effort to enforce submission to an ill-considered
and arbitrary policy on the part of the king and his ministers. The
feeling by which the king’s party was guided, in the treatment of the
American question, was very much the same as the feeling which
lately inspired the Tory criticisms upon Gladstone’s policy in South
Africa. Lord Beaconsfield, a man in some respects not Mr.
unlike Charles Townshend, bequeathed to his Gladstone
successor a miserable quarrel with the Dutch farmers and the
of the Transvaal; and Mr. Gladstone, after examining Boers
the case on its merits, had the moral courage to
acknowledge that England was wrong, and to concede the demands
of the Boers, even after serious military defeat at their hands.
Perhaps no other public act of England in the nineteenth century has
done her greater honour than this. But said the Jingoes, All the world
will now laugh at Englishmen, and call them cowards. In order to
vindicate the military prestige of England, the true policy would be,
forsooth, to prolong the war until the Boers had been once
thoroughly defeated, and then acknowledge the soundness of their
position. Just as if the whole world did not know, as well as it can
possibly know anything, that whatever qualities the English nation
may lack, it certainly does not lack courage, or the ability to win
victories in a good cause! All honour to the Christian statesman who
dares to leave England’s military prestige to be vindicated by the
glorious records of a thousand years, and even in the hour of well-
merited defeat sets a higher value on political justice than on a
reputation for dealing hard blows! Such incidents as this are big with
hope for the future. They show us what sort of political morality our
children’s children may expect to see, when mankind shall have come
somewhat nearer toward being truly civilized.
In the eighteenth century, no such exhibition of good sense and
good feeling, in the interest of political justice, could have been
expected from any European statesman, unless from a Turgot or a
Chatham. But Charles Townshend was not even called upon to
exercise any such self-control. Had he simply taken Alderman
Beckford’s advice, and done nothing, all would have been well; but
his meddling had now put the government into a position which it
was ruinous to maintain, but from which it was difficult to retreat.
American tradition rightly lays the chief blame for the troubles which
brought on the Revolutionary War to George III.; but, in fairness, it is
well to remember that he did not suggest Townshend’s measures,
though he zealously adopted and cherished them when once
propounded. The blame for wantonly throwing the apple of discord
belongs to Townshend more than to any one else.
Death of
After doing this, within three months from the time his Townshend
bill had passed the House of Commons, Townshend
was seized with a fever and died at the age of forty-one. A man of
extraordinary gifts, but without a trace of earnest moral conviction,
he had entered upon a splendid career; but his insincere nature,
which turned everything into jest, had stamped itself upon his work.
He bequeathed to his country nothing but the quarrel which was
soon to deprive her of the grandest part of that empire upon which
the sun shall never set.

GEORGE III
If Townshend’s immediate object in originating these measures
was to curry favour with George III., and get the lion’s share in the
disposal of the king’s ample corruption-fund, he had doubtless gone
to work in the right way. The king was delighted with
His political
Townshend’s measures, and after the sudden death of legacy to
his minister he made them his own, and staked his George III.
whole political career as a monarch upon their success.
These measures were the fatal legacy which the brighter political
charlatan left to the duller political fanatic. The fierce persistency with
which George now sought to force Townshend’s measures upon the
Americans partook of the nature of fanaticism, and we shall not
understand it unless we bear in mind the state of political parties in
England between 1760 and 1784. When George III. came to the
throne, in 1760, England had been governed for more than half a
century by the great Whig families which had been brought into the
foreground by the revolution of 1688. The Tories had been utterly
discredited and cast out of political life by reason of their willingness
to conspire with the Stuart pretenders in disturbing the peace of the
country. Cabinet government, in its modern form, had begun to grow
up during the long and prosperous administration of Sir Robert
Walpole, who was the first English prime minister in the full sense.
Under Walpole’s wise and powerful sway, the first two Georges had
possessed scarcely more than the shadow of sovereignty. It was the
third George’s ambition to become a real king, like the king of France
or the king of Spain. From earliest babyhood, his mother had forever
been impressing upon him the precept, ”George, be king!” and this
simple lesson had constituted pretty much the whole of his
education. Popular tradition regards him as the most Character of
ignorant king that ever sat upon the English throne; George III.
and so far as general culture is concerned, this opinion
is undoubtedly correct. He used to wonder what people could find to
admire in such a wretched driveller as Shakespeare, and he never
was capable of understanding any problem which required the
slightest trace of imagination or of generalizing power. Nevertheless,
the popular American tradition undoubtedly errs in exaggerating his
stupidity and laying too little stress upon the worst side of his
character. George III. was not destitute of a certain kind of ability,
which often gets highly rated in this not too clear-sighted world. He
could see an immediate end very distinctly, and acquired considerable
power from the dogged industry with which he pursued it. In an age
when some of the noblest English statesmen
drank their gallon of strong wine daily, or sat
late at the gambling-table, or lived in scarcely
hidden concubinage, George III. was
decorous in personal habits and pure in
domestic relations, and no banker’s clerk in London applied himself to
the details of business more industriously than he. He had a genuine
talent for administration, and he devoted this talent most assiduously
to selfish ends. Scantily endowed with human sympathy, and almost
boorishly stiff in his ordinary unstudied manner, he could be smooth
as oil whenever he liked. He was an adept in gaining men’s
confidence by a show of interest, and securing their aid by dint of fair
promises; and when he found them of no further use, he could turn
them adrift with wanton insult. Any one who dared to disagree with
him upon even the slightest point of policy he straightway regarded
as a natural enemy, and pursued him ever afterward with vindictive
hatred. As a natural consequence, he surrounded himself with weak
and short-sighted advisers, and toward all statesmen of broad views
and independent character he nursed the bitterest rancour. He had
little faith in human honour or rectitude, and in pursuing an end he
was seldom deterred by scruples.
Such was the man who, on coming to the throne in 1760, had it
for his first and chiefest thought to break down the growing system
of cabinet government in England. For the moment circumstances
seemed to favour him. The ascendancy of the great Whig families
was endangered on two sides. On the one hand, the Tory party had
outlived that idle, romantic love for the Stuarts upon which it found it
impossible to thrive. The Tories began coming to court
English
again, and they gave the new king all the benefit of parties
their superstitious theories of high prerogative and between
divine right. On the other hand, a strong popular 1760 and
feeling was beginning to grow up against 1784
parliamentary government as conducted by the old
Whig families. The House of Commons no longer fairly represented
the people. Ancient boroughs, which possessed but a handful of
population, or, like Old Sarum, had no inhabitants at all, still sent
their representatives to Parliament, while great cities of recent
growth, such as Birmingham and Leeds, were unrepresented. To a
great extent, it was the most progressive parts of the kingdom which
were thus excluded from a share in the government, while the rotten
boroughs were disposed of by secret lobbying, or even by open
bargain and sale. A few Whig families, the heads of which sat in the
House of Lords, thus virtually owned a considerable part of the House
of Commons; and, under such circumstances, it was not at all strange
that Parliament should sometimes, as in the Wilkes case, array itself
in flat opposition to the will of the people. The only wonder is that
there were not more such scandals. The party of “Old Whigs,”
numbering in its ranks some of the ablest and most patriotic men in
England, was contented with this state of things, upon which it had
thrived for two generations, and could not be made to understand
the iniquity of it,—any more than an old cut-and-dried American
politician in our time can be made to understand the iniquity of the
“spoils system.” Of this party the Marquis of Rockingham was the
political leader, and Edmund Burke was the great representative
statesman. In strong opposition to the Old Whig policy there had
grown up the party of New Whigs, bent upon bringing about some
measure of parliamentary reform, whereby the House of Commons
might truly represent the people of Great Britain. In Parliament this
party was small in numbers, but weighty in character, and at its head
was the greatest Englishman of the eighteenth century, the elder
William Pitt, under whose guidance England had won her Indian
empire and established her dominion over the seas, while she had
driven the French from America, and enabled Frederick the Great to
lay the foundations of modern Germany.
 Now when George III. came to the throne, he took
George III.
advantage of this division in the two parties in order to as a
break down the power of the Old Whig families, which politician
so long had ruled the country. To this end he used the
revived Tory party with great effect, and bid against the Old Whigs
for the rotten boroughs; and in playing off one set of prejudices and
interests against another, he displayed in the highest degree the
cunning and craft of a self-seeking politician. His ordinary methods
would have aroused the envy of Tammany. While engaged in such
work, he had sense enough to see that the party from which he had
most to fear was that of the New Whigs, whose scheme of
parliamentary reform, if ever successful, would deprive him of the
machinery of corruption upon which he relied. Much as he hated the
Old Whig families, he hated Pitt and his followers still more heartily.
He was perpetually denouncing Pitt as a “trumpeter of sedition,” and
often vehemently declared in public, and in the most offensive
manner, that he wished that great man were dead. Such had been
his eagerness to cast discredit upon Pitt’s policy that he had utterly
lost sight of the imperial interests of England, which indeed his
narrow intelligence was incapable of comprehending. One of the first
acts of his reign had been to throw away Cuba and the Philippine
Islands, which Pitt had just conquered from Spain; while at the same
time, by leaving Prussia in the lurch before the Seven Years’ War had
fairly closed, he converted the great Frederick from one of England’s
warmest friends into one of her bitterest enemies.
This political attitude of George III. toward the
His chief
Whigs in general, and toward Pitt in particular, explains reason for
the fierce obstinacy with which he took up and carried quarrelling
on Townshend’s quarrel with the American colonies. with the
For if the American position, that there should be no Americans
taxation without representation, were once to be
granted, then it would straightway become necessary to admit the
principles of parliamentary reform. The same principle that applied to
such commonwealths as Massachusetts and Virginia would be
forthwith applied to such towns as Birmingham and Leeds. The
system of rotten boroughs would be swept away; the chief engine of
kingly corruption would thus be destroyed; a reformed House of
Commons, with the people at its back, would curb forever the
pretensions of the Crown; and the detested Lord Chatham would
become the real ruler of a renovated England, in which George III.
would be a personage of very little political importance.
In these considerations we find the explanation of the acts of
George III. which brought on the American Revolution, and we see
why it is historically correct to regard him as the person chiefly
responsible for the quarrel. The obstinacy with which he refused to
listen to a word of reason from America was largely due to the
exigencies of the political situation in which he found himself. For
him, as well as for the colonies, it was a desperate struggle for
political existence. He was glad to force on the issue in America
rather than in England, because it would be comparatively easy to
enlist British local feeling against the Americans as a remote set of
“rebels,” with whom Englishmen had no interests in common, and
thus obscure the real nature of the issue. Herein he showed himself a
cunning politician, though an ignoble statesman. By playing off
against each other the two sections of the Whig party, he continued
for a while to carry his point; and had he succeeded in overcoming
the American resistance and calling into England a well-trained army
of victorious mercenaries, the political quarrel there could hardly have
failed to develop into a civil war. A new rebellion would perhaps have
overthrown George III. as James II. had been overthrown a century
before. As it was, the victory of the Americans put an end to the
personal government of the king in 1784, so quietly that the people
scarcely realized the change.[1] A peaceful election accomplished
what otherwise could hardly have been effected without bloodshed.
So while George III. lost the fairest portion of the British Empire, it
was the sturdy Americans who, fighting the battle of freedom at once
for the Old World and for the New, ended by overwhelming his paltry
schemes for personal aggrandizement in hopeless ruin, leaving him
for posterity to contemplate as one of the most instructive examples
of short-sighted folly that modern history affords.
CHAPTER II
THE CRISIS
 Townshend was succeeded in the
exchequer by Lord North, eldest son of
the Earl of Guildford, a young man of
sound judgment, wide knowledge, and
rare sweetness of temper, but wholly
lacking in sympathy with popular
government. As leader of the House of
Commons, he was sufficiently able in
debate to hold his ground against the
fiercest attacks of Burke and Fox, but
he had no strength of will. His lazy
good-nature and his Tory principles
made him a great favourite with the
king, who, through his influence over
LORD NORTH Lord North, began now to exercise the
power of a cabinet minister, and to
take a more important part than hitherto in the direction of affairs.
Soon after North entered the cabinet, colonial affairs were taken from
Lord Shelburne and put in charge of Lord Hillsborough, a man after
the king’s own heart. Conway was dismissed from the cabinet, and
his place was taken by Lord Weymouth, who had voted against the
repeal of the Stamp Act. The Earl of Sandwich, who never spoke of
the Americans but in terms of abuse, was at the same time made
postmaster-general; and in the following year Lord Chatham resigned
the privy seal.
While the ministry, by these important changes, was becoming
more and more hostile to the just claims of the
Americans, those claims were powerfully urged
in America, both in popular literature and in
well-considered state papers. John Dickinson, at
once a devoted friend of England and an ardent
American patriot, published his celebrated Farmer’s Letters, which
were greatly admired in both countries for their temperateness of
tone and elegance of expression. In these letters,
John
Dickinson held a position quite similar to that occupied
by Burke. Recognizing that the constitutional relations Dickinson
of the colonies to the mother-country had always been
extremely vague and ill-defined, he urged that the same state of
things be kept up forever through a genuine English feeling of
compromise, which should refrain from pushing any abstract theory
of sovereignty to its extreme logical conclusions. At the same time,
he declared that the Townshend revenue acts were “a most
dangerous innovation” upon the liberties of the people, and
significantly hinted, that, should the ministry persevere in its
tyrannical policy, “English history affords examples of resistance by
force.”
While Dickinson was publishing these letters,
The
Samuel Adams wrote for the Massachusetts assembly a Massachuse
series of addresses to the ministry, a petition to the tts circular
king, and a circular letter to the assemblies of the letter
other colonies. In these very able state papers, Adams
declared that a proper representation of American interests in the
British Parliament was impracticable, and that, in accordance with the
spirit of the English Constitution, no taxes could be levied in America
except by the colonial legislatures. He argued that the Townshend
acts were unconstitutional, and asked that they should be repealed,
and that the colonies should resume the position which they had
occupied before the beginning of the present troubles. The petition to
the king was couched in beautiful and
touching language, but the author seems
to have understood very well how little
effect it was likely to produce. His
daughter, Mrs. Wells, used to tell how
one evening, as her father had just
finished writing this petition, and had
taken up his hat to go out, she observed
that the paper would soon be touched by
the royal hand. “More likely, my dear,” he
replied, “it will be spurned by the royal
foot!” Adams rightly expected much more
from the circular letter to the other colonies, in which he invited them
to coöperate with Massachusetts in resisting the Townshend acts, and
in petitioning for their repeal. The assembly, having adopted all these
papers by a large majority, was forthwith prorogued by Governor
Bernard, who, in a violent speech, called them demagogues to whose
happiness “everlasting contention was necessary.” But the work was
done. The circular letter brought encouraging replies from the other
colonies. The condemnation of the Townshend acts was unanimous,
and leading merchants in most of the towns entered into agreements
not to import any more English goods until the acts should be
repealed. Ladies formed associations, under the name of Daughters
of Liberty, pledging themselves to wear homespun clothes and to
abstain from drinking tea. The feeling of the country was thus plainly
enough expressed, but nowhere as yet was there any riot or disorder,
and no one as yet, except, perhaps, Samuel Adams, had begun to
think of a political separation from England. Even he did not look
upon such a course as desirable, but the treatment of his
remonstrances by the king and the ministry soon led him to change
his opinion.
 The petition of the Massachusetts assembly was received by the
king with silent contempt, but the circular letter threw him into a
rage. In cabinet meeting, it was pronounced to be little better than
an overt act of rebellion, and the ministers were encouraged in this
opinion by letters from Bernard, who represented the whole affair as
the wicked attempt of a few vile demagogues to sow the seeds of
dissension broadcast over the continent. We have before had
occasion to observe the extreme jealousy with which the Crown had
always regarded any attempt at concerted action among the colonies
which did not originate with itself. But here was an
Lord
attempt at concerted action in flagrant opposition to Hillsborough
the royal will. Lord Hillsborough instructed Bernard to ’s
command the assembly to rescind their circular letter,
and, in case of their refusal, to send them home about instructions
their business. This was to be repeated year after year, to Bernard
so that, until Massachusetts should see fit to declare
herself humbled and penitent, she must go without a legislature. At
the same time, Hillsborough ordered the assemblies in all the other
colonies to treat the Massachusetts circular with contempt,—and this,
too, under penalty of instant dissolution. From a constitutional point
of view, these arrogant orders deserve to be ranked among the
curiosities of political history. They serve to mark the rapid progress
the ministry was making in the art of misgovernment. A year before,
Townshend had suspended the New York legislature by an act of
Parliament. Now, a secretary of state, by a simple royal order,
threatened to suspend all the legislative bodies of America unless
they should vote according to his dictation.
When Hillsborough’s orders were laid before the Massachusetts
assembly, they were greeted with scorn. “We are asked to rescind,”
said Otis. “Let Britain rescind her measures, or the colonies are lost
to her forever.” Nevertheless, it was only after nine
The
days of discussion that the question was put, when the “Illustrious
assembly decided, by a vote of ninety-two to Ninety-Two”
seventeen, that it would not rescind its circular letter.
Bernard immediately dissolved the assembly, but its vote was hailed
with delight throughout the country, and the “Illustrious Ninety-Two”
became the favourite toast on all convivial occasions. Nor were the
other colonial assemblies at all readier than that of Massachusetts to
yield to the secretary’s dictation. They all expressed the most cordial
sympathy with the recommendations of the circular letter; and in
several instances they were dissolved by the governors, according to
Hillsborough’s instructions.
 FANEUIL HALL, THE CRADLE OF LIBERTY
While these fruitless remonstrances against the Townshend acts
had been preparing, the commissioners of the customs, in enforcing
the acts, had not taken sufficient pains to avoid irritating the people.
In the spring of 1768, the fifty-gun frigate Romney had
Impressmen
been sent to mount guard in the harbour of Boston, t of citizens
and while she lay there several of the citizens were
seized and impressed as seamen,—a lawless practice long afterward
common in the British navy, but already stigmatized as barbarous by
public opinion in America. As long ago as 1747, when the relations
between the colonies and the home government were quite
harmonious, resistance to the press-gang had resulted in a riot in the
streets of Boston. Now while the town was very indignant over this
lawless kidnapping of its citizens, on the 10th of June, 1768, John
Hancock’s sloop Liberty was seized at the wharf by a boat’s crew
from the Romney, for an alleged violation of the revenue laws,
though without official warrant. Insults and recriminations ensued
between the officers and the citizens assembled on the wharf, until
after a while the excitement grew into a mild form of riot, in which a
few windows were broken, some of the officers were pelted, and
finally a pleasure boat, belonging to the collector, was pulled up out
of the water, carried to the Common, and burned there, when
Hancock and Adams, arriving upon the scene, put a stop to the
commotion. A few days afterward, a town meeting was held in
Faneuil Hall; but as the crowd was too great to be contained in the
building, it was adjourned to the Old South Meeting-House, where
Otis addressed the people from the pulpit. A petition to the governor
was prepared, in which it was set forth that the impressment of
peaceful citizens was an illegal act, and that the state of the town
was as if war had been declared against it; and the governor was
requested to order the instant removal of the frigate from the
harbour. A committee of twenty-one leading citizens was appointed to
deliver this petition to the governor at his house in Jamaica Plain. In
his letters to the secretary of state Bernard professed to live in
constant fear of assassination, and was always begging for troops to
protect him against the incendiary and blackguard mob of Boston. Yet
as he looked down the beautiful road from his open window, that
summer afternoon, what he saw was not a ragged mob, armed with
knives and bludgeons, shouting “Liberty, or death!” and bearing the
head of a revenue collector aloft on the point of a pike, but a quiet
procession of eleven chaises, from which there alighted at his door
twenty-one gentlemen, as sedate and stately in demeanour as those
old Roman senators at whom the Gaulish chief so marvelled. There
followed a very affable interview, during which wine was passed
around. The next day the governor’s answer was read in town
meeting, declining to remove the frigate, but promising that in future
there should be no impressment of Massachusetts citizens; and with
this compromise the wrath of the people was for a moment
assuaged.
 Affairs of this sort, reported with gross exaggeration by the
governor and revenue commissioners to the ministry, produced in
England the impression that Boston was a lawless and riotous town,
full of cutthroats and blacklegs, whose violence could be held in
check only by martial law. Of all the misconceptions of America by
England which brought about the American Revolution, perhaps this
notion of the turbulence of Boston was the most ludicrous. During
the ten years of excitement which preceded the War of
Independence there was one disgraceful riot in Boston,—that in
which Hutchinson’s house was sacked; but in all this time not a drop
of blood was shed by the people, nor was anybody’s life for a
moment in danger at their hands. The episode of the sloop Liberty, as
here described, was a fair sample of the disorders which occurred at
Boston at periods of extreme excitement; and in any European town
in the eighteenth century it would hardly have been deemed worthy
of mention.
 Even before the affair of the Liberty, the government had made
up its mind to send troops to Boston, in order to overawe the popular
party and show them that the king and Lord Hillsborough were in
earnest. The news of the Liberty affair, however, served to remove
any hesitation that might hitherto have been felt.
Statute of
Vengeance was denounced against the insolent town Henry VIII.
of Boston. The most seditious spirits, such as Otis and concerning
Adams, must be made an example of, and thus the “treason
others might be frightened into submission. With such committed
intent, Lord Hillsborough sent over to inquire “if any abroad”
person had committed any acts which, under the statutes of Henry
VIII. against treason committed abroad, might justify their being
brought to England for trial.” This raking-up of an obsolete statute,
enacted at one of the worst periods of English history, and before
England had any colonies at all, was extremely injudicious. But
besides all this, continued Hillsborough, the town meeting, that
nursery of sedition, must be put down or overawed; and in
pursuance of this scheme, two regiments of soldiers and a frigate
were to be sent over to Boston at the ministry’s earliest convenience.
To make an example of Boston, it was thought, would have a
wholesome effect upon the temper of the Americans.
 LANDING OF THE TROOPS IN BOSTON, 1768

CASTLE WILLIAM, BOSTON HARBOUR

It was now, in the summer of 1768, that Samuel Adams made up
his mind that there was no hope of redress from the British
government, and that the only remedy was to be found in the
assertion of political independence by the American colonies. The
courteous petitions and temperate remonstrances of the American
assemblies had been met, not by rational arguments,
Samuel
but by insulting and illegal royal orders; and now at Adams
last an army was on the way from England to enforce makes up
the tyrannical measures of government, and to terrify his mind,
the people into submission. Accordingly, Adams came 1768
to the conclusion that the only proper course for the
colonies was to declare themselves independent of Great Britain, to
unite together in a permanent confederation, and to invite European
alliances. We have his own word for the fact that from this moment
until the Declaration of Independence, in 1776, he consecrated all his
energies, with burning enthusiasm, upon the attainment of that great
object. Yet in 1768 no one knew better than Samuel Adams that the
time had not yet come when his bold policy could be safely adopted,
and that any premature attempt at armed resistance on the part of
Massachusetts might prove fatal. At this time, probably no other
American statesman had thought the matter out so far as to reach
Adams’s conclusions. No American had as yet felt any desire to
terminate the political connection with England. Even those who most
thoroughly condemned the measures of the government did not
consider the case hopeless, but believed that in one way or another a
peaceful solution was still attainable. For a long time this attitude was
sincerely and patiently maintained. Even Washington, when he came
to take command of the army at Cambridge, after the battle of
Bunker Hill, had not made up his mind that the object of the war was
to be the independence of the colonies. In the same month of July,
1775, Jefferson said expressly, “We have not raised armies with
designs of separating from Great Britain and establishing independent
states. Necessity has not yet driven us into that desperate measure.”
The Declaration of Independence was at last brought about only with
difficulty and after prolonged discussion. Our great-great-
grandfathers looked upon themselves as Englishmen, and felt proud
of their connection with England. Their determination to resist
arbitrary measures was at first in no way associated in their minds
with disaffection toward the mother-country. Besides this, the task of
effecting a separation by military measures seemed to most persons
quite hopeless. It was not until after Bunker Hill had shown that
American soldiers were a match for British soldiers in the field, and
after Washington’s capture of Boston had shown that the enemy
really could be dislodged from a whole section of the country, that
the more hopeful patriots began to feel confident of the ultimate
success of a war for independence. It is hard for us now to realize
how terrible the difficulties seemed to the men who surmounted
them. Throughout the war, beside the Tories who openly sympathized
with the enemy, there were many worthy people who thought we
were “going too far,” and who magnified our losses and depreciated
our gains,—quite like the people who, in the War of Secession, used
to be called “croakers.” The depression of even the boldest, after
such defeats as that of Long Island, was dreadful. How inadequate
was the general sense of our real strength, how dim the general
comprehension of the great events that were happening, may best
be seen in the satirical writings of some of the loyalists. At the time
of the French alliance, there were many who predicted that the result
of this step would be to undo the work of the Seven Years’ War, to
reinstate the French in America with full control over the thirteen
colonies, and to establish despotism and popery all over the
continent. A satirical pamphlet, published in 1779, just ten years
before the Bastille was torn down in Paris, drew an imaginary picture
of a Bastille which ten years later was to stand in New York, and,
with still further license of fantasy, portrayed Samuel Adams in the
garb of a Dominican friar. Such nonsense is of course no index to the
sentiments or the beliefs of the patriotic American people, but the
mere fact that it could occur to anybody shows how hard it was for
people to realize how competent America was to take care of herself.
The more we reflect upon the slowness with which the country came
to the full consciousness of its power and importance, the more fully
we bring ourselves to realize how unwilling America was to tear
herself asunder from England, and how the Declaration of
Independence was only at last resorted to when it had become
evident that no other course was compatible with the preservation of
our self-respect; the more thoroughly we realize all this, the nearer
we shall come toward duly estimating the fact that in 1768, seven
years before the battle of Lexington, the master mind of Samuel
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