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- Herausgeber: John Wiley & Sons
- Kategorie: Wissenschaft und neue Technologien
- Serie: Plant Breeding Reviews
- Sprache: Englisch
Plant Breeding Reviews presents state-of-the-art reviews on plant genetics and the breeding of all types of crops by both traditional means and molecular methods. Many of the crops widely grown today stem from a very narrow genetic base; understanding and preserving crop genetic resources is vital to the security of food systems worldwide. The emphasis of the series is on methodology, a fundamental understanding of crop genetics, and applications to major crops.
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Veröffentlichungsjahr: 2018
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Table of Contents
Cover
Title Page
Contributors
1 Hari Deo Upadhyaya
ABBREVIATIONS
I. INTRODUCTION
II. BIOGRAPHICAL SKETCH
III. CONTRIBUTIONS
IV. UPADHYAYA, THE MAN
V. PUBLICATIONS
VI. PRODUCTS
REFERENCES CITED AND FURTHER READING
2 Crop Improvement Using Genome Editing
ABBREVIATIONS
I. INTRODUCTION
II. CONCEPTUAL FRAMEWORK FOR GENOME EDITING
III. PLANT TRANSFORMATION STRATEGIES
IV. HARNESSING BREAKS FOR TARGETED MUTAGENESIS
V. PRECISION GENE EDITING VIA HOMOLOGOUS RECOMBINATION
VI. GENOME EDITING AT THE GENOME LEVEL
VII. FUTURE PERSPECTIVES
ACKNOWLEDGEMENTS
LITERATURE CITED
3 Development and Commercialization of CMS Pigeonpea Hybrids
ABBREVIATIONS
I. INTRODUCTION
II. REPRODUCTIVE CYCLE AND MORPHOLOGY OF PIGEONPEA
III. CROP PRODUCTION
IV. EXTENT AND NATURE OF HETEROSIS IN PIGEONPEA
V. GENETIC MALE STERILITY‐BASED HYBRID TECHNOLOGY
VI. TEMPERATURE‐SENSITIVE MALE STERILITY
VII. CYTOPLASMIC‐NUCLEAR MALE STERILITY‐BASED HYBRID TECHNOLOGY
VIII. BREEDING NEW HYBRID PARENTS
IX. APPLICATION OF GENOMICS IN BREEDING HYBRIDS
X. COMMERCIALIZATION OF HYBRID PIGEONPEA TECHNOLOGY
XI. OUTLOOK
ACKNOWLEDGEMENTS
LITERATURE CITED
4 The Evolution of Potato Breeding
ABBREVIATIONS
I. INTRODUCTION
II. CLASSIFICATION OF CULTIVATED POTATO
III. ORIGIN OF THE CULTIVATED POTATO
IV. DYNAMICS OF POTATO LANDRACE EVOLUTION
V. ORIGIN OF THE EUROPEAN POTATO
VI. NINETEENTH CENTURY POTATO BREEDING
VII. EARLY TWENTIETH CENTURY POTATO BREEDING
VIII. CONVENTIONAL POTATO BREEDING
IX. LATE TWENTIETH CENTURY POTATO BREEDING
X. TWENTY‐FIRST CENTURY POTATO BREEDING
XI. CONCLUSIONS
LITERATURE CITED
5 Flavour Evaluation for Plant Breeders
ABBREVIATIONS
I. INTRODUCTION
II. TYPES OF RAPID SENSORY ANALYSIS METHODS
III. DATA ANALYSIS FOR RAPID SENSORY METHODS
IV. EXAMPLE OF USING SENSORY ANALYSIS FOR BREEDING
V. OUTLOOK
ACKNOWLEDGEMENTS
LITERATURE CITED
6 The Genetic Improvement of Black Walnut for Timber Production
ABBREVIATIONS
I. INTRODUCTION
II. BIOLOGY OF BLACK WALNUT
III. BREEDING
IV. EVALUATION OF HERITABLE TRAITS
V. HOST PLANT RESISTANCE TO PATHOGENS AND INSECT PESTS
VI. PROPAGATION
VII. PLOT MANAGEMENT
VIII. FUTURE DIRECTIONS
LITERATURE CITED
7 A Life in Horticulture and Plant Breeding
ABBREVIATIONS
I. INTRODUCTION
II. HONORS AND COMMENDATIONS
III. STUDENTS AND TEACHING
IV. EDITORIAL WORK
V. BOOKS AND PROCEEDINGS
VI. RESEARCH
VII. PUBLIC ADDRESSES, INVITED SEMINARS AND SPEECHES
VIII. SERVICE CONTRIBUTIONS
IX. EPILOGUE
LITERATURE CITED
Auhtor Index
Subject Index
Cumulative Subject Index
Cumulative Contributor Index
End User License Agreement
List of Tables
Chapter 01
Table 1.1. Core collections formed by Hari Deo Upadhyaya in chickpea, groundnut, pearl millet, pigeonpea, and small millets.
Table 1.2. Mini‐core collections formed by Hari Deo Upadhyaya in chickpea, groundnut, pearl millet, pigeonpea, sorghum, and small millet.
Table 1.3. Composite collections and reference sets formed by Hari Deo Upadhyaya in chickpea, groundnut, pearl millet, pigeonpea, sorghum, finger millet, and foxtail millet.
Table 1.4. Some of the awards and honours received by Hari Deo Upadhyaya from 2002 to 2017.
Table 1.5. List of 27 groundnut cultivars and their local names released in 18 countries in Africa and Asia.
Chapter 02
Table 2.1. Applications of genome editing.
Table 2.2. Sequence‐specific nucleases used for genome editing.
Table 2.3. Plant transformation systems used for genome editing.
Chapter 03
Table 3.1. Area, production, and grain yield of the main pigeonpea growing countries.
Table 3.2. Pigeonpea maturity groups based on days to flowering and maturity of genotypes planted by mid‐June at ICRISAT in Patancheru, India (17°N 30’, 78°16’E, 545 m.a.s.l.) Table based on Vales
et al
. (Vales et al., 2012) and Green
et al
. (Green et al., 1979). Expanded to show divisions based on days to 75% maturity.
Table 3.3. Distinctive characteristics of genetic male sterility (GMS) systems reported over time in pigeonpea.
Table 3.4. Summary of standard heterosis
z
(%) reported in genetic male sterile‐based experimental hybrids produced at ICRISAT in the early phases.
Table 3.5. Standard heterosis of genetic male sterile pigeonpea hybrids belonging to early and medium maturity, released across India in the 1990s.
Table 3.6. Number of sterile/fertile plants recorded in temperature‐sensitive male sterile (TSMS) selections planted under an insect‐proof net at ICRISAT, Patancheru (17°N), India. The plants were male‐sterile at temperatures above 25°C (could be used to produce hybrids). The same plants became fully fertile at temperatures below 24°C (produced self‐pollinated seed of the maternal line).
Table 3.7. Cytoplasmic male sterility (CMS) systems reported in pigeonpea, based on various cytoplasm donors.
Table 3.8. Phenotypic variation observed for important traits among pigeonpea fertility restorers (R lines) and maintainers (B lines) used to produce cytoplasmic male sterile (CMS) hybrids.
Table 3.9. Performance of medium maturity pigeonpea hybrids involving A‐lines with obcordate leaf shape.
Table 3.10. Distribution of CMS A lines and restorer lines in different heterotic groups based on diversity using SSR markers.
Table 3.11. Yield and standard heterosis of CMS pigeonpea hybrids, representing three major maturity groups, recorded in multi‐location trials across India.
Table 3.12. Yield and standard heterosis of the cytoplasmic male sterile medium maturity pigeonpea hybrid ICPH 2671 in comparison with cultivar Maruti, planted across India in multi‐location trials from 2005–2008.
Table 3.13. Yield of medium maturity cytoplasmic male sterile pigeonpea hybrid ICPH 2671 and popular cultivar Maruti recorded from on‐farm trials spread over four states in India in 2008 and 2009.
Table 3.14. Yield and standard heterosis of medium maturity cytoplasmic male sterile pigeonpea hybrid ICPH 3762 and popular cultivar Asha obtained from on‐farm trials conducted in four districts of Odisha, India, in 2013.
Table 3.15. Yield and standard heterosis of medium maturity cytoplasmic male sterile pigeonpea hybrid ICPH 2740 and popular cultivar Asha obtained from on‐farm trials conducted in four states in India from 2009 to 2011.
Table 3.16. Main areas in India suited for cytoplasmic male sterile pigeonpea hybrid seed production (A × R) based on accomplished yields in farmers’ fields, 2008–09.
Table 3.17. Better parent heterosis for yield, male sterility systems, and out‐crossing rates recorded in several food legumes.
Chapter 04
Table 4.1. Typical schedule for a conventional potato breeding programme.
Chapter 05
Table 5.1. Production and culinary traits of interest to farmers for the 12 species included in the 2014 and 2015 variety trials.
Table 5.2. ANOVA of crew flavour evaluation on tomatoes in 2016 using intensity scaling.
Chapter 07
Table 7.1. Honours and Commendations of Professor Jules Janick.
Table 7.2. Postdoctoral associates, graduate students, and theses supervision.
Table 7.3. Journal volumes and reviews where Jules Janick has served as editor.
Table 7.4. Books and proceedings written and edited by Jules Janick.
Table 7.5. Patents, plant patents, and plant variety protection certificates of Jules Janick.
List of Illustrations
Chapter 01
Figure 1.1.
Cajanus albicans
, a wild species from a secondary genepool with many desirable characteristics, and a potential source for gene introgression in cultivated pigeonpea.
Figure 1.2. Cryptic genetic variation captured in groundnut by Hari Deo Upadhyaya that originated from a cross between TMV 2 (a leading groundnut variety in southern India) and amphidiploid (TxAG 6).
Figure 1.3. ICGV 91114, an improved groundnut variety that was developed by Hari Deo Upadhyaya and shows early maturity (90 days), drought tolerance and resistance to rust and late leaf spot, which is now popularly grown in India.
Chapter 02
Figure 2.1. Sequence‐specific nuclease (SSN) platforms and variations used for genome editing. Homing endonucleases (meganucleases) are natural endonucleases with multifunctional domains, capable of binding double‐strand DNA targets and making double‐stranded breaks (DSBs; paired scissors). Zinc finger nucleases (ZFNs) are synthetic nucleases composed of assembled zinc finger DNA binding domains (black hexagons) and the FokI nuclease domain (grey trapezoids). Transcription activator‐like effector nucleases (TALENs) are synthetic nucleases composed of assembled TAL effector DNA binding domains (black boxes) and the FokI nuclease. Clustered regularly interspaced short palindromic repeats/CRISPR‐associated system (CRISPR/Cas) is a RNA‐guided endonuclease system employing the Cas9 nuclease (grey circle), with RuvC and HNH nuclease domains and a single‐guide RNA to guide Cas9 to DNA targets. Modification of Cas9 where RuvC or HNH nuclease domain activity is eliminated results in creation of D10A and H840A CRISPR/Cas nickases, respectively capable of producing single‐strand breaks (SSBs; single scissors). Replacement of the TALEN FokI nuclease domain with the I‐Tev nuclease domain results a compact TALEN (cTALEN) nickase capable of creating SSBs. Target DNA recognition lengths are in base pairs (bp).
Figure 2.2. DNA damage and DNA repair pathways used for genome editing. DNA damage, in the form of double‐strand breaks (DSBs) and single‐strand breaks (SSBs or ‘nicks’), underlie most applications of genome editing. DNA damage can originate both from natural (ionizing radiation, chemical, cellular processes, and oxidative damage) and synthetic (sequence‐specific nucleases and ‘nickases’) sources, and typically results in random or directed DNA damage, respectively. DSBs are a severe form of DNA damage, which induce a strong DNA repair response, primarily of the non‐homologous end‐joining (NHEJ) pathway (thick arrow). SSBs, or ‘nicks’, are a less severe form of DNA damage, but can stimulate the homologous recombination (HR) pathway (thin arrow), or can be directed to an adjacent region to induce DSBs and NHEJ. NHEJ follows two sub‐pathways, called canonically NHEJ (cNHEJ) or alternative NHEJ (aNHEJ), for low‐fidelity repair and loss or gain of sequence at the break site and frame‐shift mutations (Targeted mutagenesis). A sub‐pathway of HR, called single‐strand annealing (SSA), can also result in loss of sequence and frame‐shift mutations in the form of targeted mutagenesis. An alternative HR sub‐pathway, called synthesis‐dependent strand annealing (SDSA), uses a DNA repair template (Repair template) to repair both SSBs and DSBs, and is capable of introducing new sequence at the break site with high fidelity (grey stars), resulting in so‐called gene targeting. Gene targeting is the most useful form of genome editing but is least preferred in most plant cells, compared with other sub‐pathways of DNA repair.
Figure 2.3. Possible modes of inheritance for SSNs and targeted modifications. In this example, a selfing crop species (line ‘Inbred A’) is stably transformed with a CRISPR/Cas9 transgene that targets a specific mutation, and the desired genotype is a homozygous mutated line with no transgene. The transgene and the target locus are unlinked, permitting independent segregation of the two loci. The subscript accompanying each letter represents the number of generations, since the introduction of the transgene (‘T’) or induction of a mutation (‘M’). Solid lines represent direct descent from the previous generation, while dashed lines represent multiple generations. Individuals with fixed states (no longer segregating) are shown in rectangles (undesired genotypes) or circles (desired genotypes). Individuals not within rectangles or circles are heterozygous for at least one locus, or still capable of generating new mutations (i.e. carrying the transgene in the heterozygous or homozygous state). The diagram does not show all possible outcomes, but instead represents several possible scenarios that illustrate the complexity of tracking the two segregating loci and recovering the desired plant.
Chapter 03
Figure 3.1. Short duration non‐determinate pigeonpea CMS hybrid ICPH 2433 at the flowering stage. ICRISAT, Telangana, India.
Figure 3.2. Medium duration non‐determinate pigeonpea CMS hybrid ICPH 2740 at podding stage. Jalgaon, Maharashtra, India.
Chapter 05
Figure 5.1. Results of the PCA on LS Mean values from crew flavour evaluation:
a
shows the factor loadings of the different flavour components on the first two principal components and
b
shows the placement of varieties. The first two principal components capture 85.81% of the total variation.
Figure 5.2. Correlation between perceived flavour intensity least squared means and overall flavour appreciation least squared means for the crew flavour evaluation. Error bars shown are standard error of the mean.
Figure 5.3. Consensus plot from MFA of chef 2015 evaluation of tomato varieties. Axes were labelled by chefs in advance of the tasting.
Figure 5.4. Consensus plot from MFA of chef 2016 evaluation of tomato varieties. Axes were labelled based on the results of the crew PCA flavour component factor loadings.
Figure 5.5. Correlation between perceived flavour intensity and overall flavour appreciation for the chef flavour evaluation. Each point corresponds to one chef’s rating of one sample.
Chapter 07
Figure 7.1. Professor Jules Janick.
Figure 7.2. Jules Janick in 1945 at Victory Farm Corps, obtaining agricultural training to enter Cornell University.
Figure 7.3. Jules Janick with ‘GoldRush’ apple.
Guide
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PLANT BREEDING REVIEWS
Volume 41
Edited by
Irwin Goldman
University of Wisconsin‐MadisonWisconsin, USA
This edition first published 2018© 2018 John Wiley & Sons, Inc.
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Library of Congress Catalog Control Number: 83‐641963
Cover Design: WileyCover Illustration: © browndogstudios/Gettyimages
Contributors
Nathaniel M Butler, Department of Horticulture, University of Wisconsin, Madison, WI
Mark V Coggeshall, USDA Forest Service, Northern Research Station, Hardwood Tree Improvement and Regeneration Center, West State Street, West Lafayette, IN, USA
JC Dawson, Department of Horticulture, University of Wisconsin‐Madison, Madison, WI
Irwin Goldman, Department of Horticulture, University of Wisconsin‐Madison, Madison, WI
GK Healy, Department of Horticulture, University of Wisconsin‐Madison, Madison, WI
Shelley H Jansky, United States Department of Agriculture – Agricultural Research Service, and University of Wisconsin, Madison, WI
Jiming Jiang, Department of Horticulture, University of Wisconsin, Madison, WI
Sangam L Dwivedi, International Crops Research Institute for the Semi‐Arid Tropics (ICRISAT), Patancheru, Telangana, India
James R McKenna, USDA Forest Service, Northern Research Station, Hardwood Tree Improvement and Regeneration Center, West State Street, West Lafayette, IN, USA
Rodomiro Ortiz, Swedish University of Agricultural Sciences, Department of Plant Breeding, Sweden
KB Saxena, International Crops Research Institute for the Semi‐Arid Tropics (ICRISAT), Patancheru, Telangana, India
D Sharma, International Crops Research Institute for the Semi‐Arid Tropics (ICRISAT), Patancheru, Telangana, India
David M Spooner, United States Department of Agriculture – Agricultural Research Service, and University of Wisconsin, Madison, WI
Robert M Stupar, Department of Agronomy and Plant Genetics, University of Minnesota, St. Paul, MN
MI Vales, Department of Horticultural Sciences, Texas A&M University, College Station, TX
1Hari Deo Upadhyaya: Plant Breeder, Geneticist and Genetic Resources Specialist
Sangam L Dwivedi
International Crops Research Institute for the Semi‐Arid Tropics (ICRISAT), Patancheru, Telangana, India
ABSTRACT
This chapter discusses Hari Deo Upadhyaya, a plant breeder, geneticist and genetic resources specialist, and his contributions in management and utilization of genetic resources, molecular biology and biometrics, and in groundnut breeding. Hari’s contributions in genetic resources include enriching germplasm collections; forming representative subsets in the form of core and/or mini‐core collections in chickpea, groundnut, pigeonpea, pearl millet, sorghum, and six small millets; unlocking population structures, diversity and association genetics; and identifying genetically diverse and agronomically desirable germplasm accessions for use in crop breeding. The Consultative Group on International Agriculture Research (CGIAR) recognized his concept and process of forming mini‐core collection as International Public Goods (IPGs) and researchers worldwide are now using mini core‐collections as useful genetic resources in breeding and genomics of the aforementioned crops. A genebank manager’s role isn’t just confined to collection, maintenance, and archiving germplasm. Hari’s spirited efforts prove so and they led many to realize the abundant opportunities to mine and enhance the value of the genetic resources in crop improvement programs. As a geneticist, his seminal work on wilt resistance in chickpea laid a strong foundation for the wilt resistance breeding programs globally. His contributions as a groundnut breeder resulted in the release of 27 cultivars in 18 countries, some widely grown, and 24 elite germplasm releases with unique characteristics made available to groundnut researchers worldwide. Hari’s inimitable ability and scientific competence allowed him to collaborate with diverse groups and institutions worldwide. His scientific contributions in germplasm research and groundnut breeding have been recognized with several prestigious global awards and honors. A prolific writer and with immense passion for teaching, Hari Upadhyaya has established a school of his own for the management, evaluation and use of genetic resources for crop improvement.
KEYWORDS: Breeding, Climate resilient germplasm, core and mini‐core collections, crop wild relatives, cultivars, elite germplasm, farmers participatory variety selection, molecular breeding, population structure and diversity, on‐farm conservation of germplasm
OUTLINE
ABBREVIATIONS
INTRODUCTION
BIOGRAPHICAL SKETCH
CONTRIBUTIONS
Genetic Resources Management and Use
Representative Subsets
Climate‐resilient Germplasm
Seed Nutrient‐dense Germplasm
Bioenergy
Germplasm Use in Breeding
On‐farm Conservation and Use of Diversity
Wild Relatives and Cultigen Genepool
Gaps in Collections
Molecular Biology and Biometrics
Population Structure and Diversity
Genome‐wide Association Mapping
Candidate Genes Associated with Agronomically Beneficial Traits
Ethnolinguistic Groups Shaped Sorghum Diversity in Africa
Genome Sequencing
Groundnut Breeding
Early Maturity
Drought Tolerance
Aflatoxin Resistance
Farmers Participatory Varietal Selection
Chickpea Breeding
UPADHYAYA, THE MAN
Personality
Educator and Leader
International Collaborations
Recognition
Awards
Honours
Service
PUBLICATIONS
PRODUCTS
Cultivars
Registrations
REFERENCES CITED AND FURTHER READING
ABBREVIATIONS
ASA
American Society of Agronomy
CGIARC
Consultative Group on International Agricultural Research Consortium
CSSA
Crop Science Society of America
ICRISAT
International Crops Research Institute for Semi‐Arid Tropics
NARS
National Agricultural Research Systems
R4D
Research for development
SNP
Single nucleotide polymorphisms
I. INTRODUCTION
Hari Deo Upadhyaya, whom many of us know as Hari, has been known to me since 1980, when he joined the International Crops Research Institute for the Semi‐Arid Tropics (ICRISAT), Patancheru, India, as a postdoctoral fellow in chickpea breeding. After completing his postdoctoral assignment at ICRISAT, Hari then moved for a short period to work as the Pool Officer at ‘GB Pant’ University of Agriculture and Technology (GBPUAT), Pantnagar, India, the first agricultural university established on a US ‘Land Grant’ pattern in India. He then took up a regular position at the University of Agriculture Sciences (UAS), Dharwad, India, where he worked for almost for eight years, first as a soybean breeder (as Assistant Professor), and then as the head of the oilseeds scheme and a groundnut breeder (as Associate Professor). He did a remarkable job as an oilseed breeder, and he set up and took the soybean and groundnut breeding programs to newer heights.
In 1991, Hari returned to ICRISAT as a Senior Groundnut Breeder. In late 1997, ICRISAT reorganized its research portfolio, and moved Hari on a part‐time basis to the Genetic Resources Unit, as part of the Crop Improvement Program. In 2002, Hari was appointed as a Principal Scientist and Head of the Genebank, ICRISAT, Patancheru, India, a position he still holds in the ‘new organizational structure’, where he has to manage the ICRISAT administrative Research for Development (R4D) portfolios with respect to management and utilization of genetic resources in crop improvement programs.
Hari knows very well that greater use of germplasm in crop breeding is the way forward for better conservation and use of genetic resources, and to address food and nutritional security in the developing world. As a principal scientist (in genetic resources), Hari performed exceedingly well, while promoting the greater use of genetic resources in crop improvement. Today, the representative subsets (i.e. the core and mini‐core collections) of the ICRISAT crops (i.e. chickpea, groundnut, pearl millet, pigeonpea, sorghum, finger millet) and small millets (i.e. barnyard millet, foxtail millet, kodo millet, little millet, proso millet) have been made available, and globally researchers are using these subsets to identify new sources of variation to support crop breeding in their respective regions.
Hari’s seminal work with Rodomiro Ortiz on the process and concept of forming the mini‐core collection has been recognized as an ‘International Public Good’. Hari has published a total of 812 articles, of which 291 have undergone international peer review. These include research articles, commissioned reviews, and book chapters, and he has averaged 11.6 such articles per year, with three articles per year as first author. Twenty‐seven cultivars of groundnut that were bred by Hari are being cultivated in 18 countries in Africa and Asia.
Over my long association with Hari, I have found him to be a person with the highest scientific competence and integrity, and a successful plant breeder and genebank manager. Hari’s leadership in managing one of the largest Consultative Group on International Agricultural Research (CGIAR) Consortium genebanks is very much reflected in a recently concluded external review, when the panel remarked that ‘The ICRISAT genebank is functioning to high technical and scientific standards, and is very good in comparison with other international genebank operations. The users of the ICRISAT genebank are satisfied and appreciation of the genebank is wide spread.’
II. BIOGRAPHICAL SKETCH
Hari was born on 12th August 1953, in the small village of Shiwala, in Khair Tehsil, District Aligarh, Uttar Pradesh, India. He is the seventh of the eight children of Mr Gopi Chand Upadhyaya and Mrs Longsri Devi Upadhyaya. He passed his high school examinations (X standard) with Biology as his main subject, and got a distinction in Mathematics. Hari did a BSc (with honours) at Aligarh Muslim University, Aligarh, India, and then moved to the GB Pant University of Agriculture and Technology, Pantnagar, India, to complete his MSc and PhD, both in Plant Breeding. Hari is married to Ms Sudha, and is blessed with two sons, Abhisheik Deo and Aaditya Deo. Interestingly, neither of his sons has followed in his footsteps, as they chose Information Technology for their career path. Hari derives great strength from his wife and children in his scientific endeavours.
III. CONTRIBUTIONS
Unlike traditional germplasm botanists and curators, whose vision is always centred on collection, conservation, characterization and documentation of germplasm, Hari’s basic training in plant breeding and genetics helped him to think beyond routine genebank activities, to include enhancing the value of genetic resources in the breeder’s perception. Plant breeders are often reluctant to use exotic germplasm, largely because of the fear of linkage drag, breakdown of co‐adapted gene complexes, and lengthening of the breeding cycle for the development of new cultivars. Hari strongly believes in promoting the use of germplasm in crop improvement programs, the generation and use of new knowledge (i.e. physiological, genetic, molecular) of trait expression and inheritance in applied breeding, and the sharing of breeding populations and advanced varieties, and also of knowledge, to help the global community to increase the production and productivity of staple food crops. Hari invested heavily to add value to the germplasm collections, and uses this in the crop breeding at ICRISAT and in the national programs globally.
A. Genetic Resources Management and Use
1. Representative Subsets.
The use of germplasm in crop improvement programs globally is restricted due to:
the large sizes of collections of many crop species;
the non‐availability of representative subsets; and
the lack of accurate and precise information on the agronomic worth of individual germplasm.
Hari saw the need, as advocated by Frankel and Brown (1984) to form reduced subsets that represent the diversity of the entire collection of a given species preserved in the genebank, and he initiated work to develop representative sets for ICRISAT mandate crops and small millets. Using passport and characterization data and statistical tools, Hari first developed the core collections (10% of the entire collection of a species stored in the genebank) for chickpea and, later, for pigeonpea, groundnut, pearl millet, and small millets (Table 1.1).
Table 1.1. Core collections formed by Hari Deo Upadhyaya in chickpea, groundnut, pearl millet, pigeonpea, and small millets.
Crop
Number of accessions used
Number of traits involved
Number of accessions in core
Reference
Pearl millet
20,766
22
2,094
Upadhyaya
et al.
, 2009a
Chickpea
16,991
13
1,956
Upadhyaya
et al.
, 2001a
Pigeonpea
12,153
14
1,290
Reddy
et al.
, 2005
Groundnut
14,310
14
1,704
Upadhyaya
et al.
, 2003
Finger millet
5,940
14
622
Upadhyaya
et al.
, 2006c
Foxtail millet
1,474
23
155
Upadhyaya
et al.
, 2008b
Proso millet
833
20
106
Upadhyaya
et al.
, 2011i
Barnyard millet
736
21
89
Upadhyaya
et al.
, 2014c
Kodo millet
656
20
75
Upadhyaya
et al.
, 2014c
Little millet
460
20
56
Upadhyaya
et al.
, 2014c
The chickpea core collection consisted of 1,956 accessions that had been selected from 16,991 accessions (Upadhyaya et al., 2001a). Rodomiro Ortiz, the then Director of Genetic Resources and the Enhancement Program, ICRISAT, challenged Hari and Paula Bramel (a co‐author with Hari) about how useful the core collections were, with such large numbers of accessions for screening a desired trait for further use in breeding. After evaluating 1,956 accessions, together with controls for one season, in an augmented design, Hari concluded that it was a Herculean task to accurately and cost‐effectively generate datasets even for the core collection accessions.
Hari and Rodomiro Ortiz discussed this and adopted the approach of re‐sampling the core collection to define a ‘core of the core’ or ‘mini‐core’, subset. Here, they used the evaluation data (22 morphological and agronomic traits) of the core collection (1956 accessions) and statistical theory to sample the variability to form the mini‐core collection (211 accessions) in chickpea. This represented the diversity that was present in the core collection, and also the entire collection, as shown by the similar means, variances, frequency distributions and preserved co‐adapted gene complexes, both for the core and mini‐core collections (Upadhyaya and Ortiz, 2001).
Hari and Rodomiro jointly wrote a manuscript on the chickpea mini core collection, with Rodomiro as corresponding author, and submitted it to Theoretical Applied Genetics. To their surprise, exactly two weeks later, they got a response from the editor to say that the manuscript was accepted for publication. This development encouraged Hari to follow this approach, and in subsequent years, he developed mini‐core collections for other crops as well (Table 1.2). In all cases, both the core and mini‐core collections fulfilled the statistical tests for the preservation of means, variances, and frequency distributions, and the co‐adapted gene complexes of the entire collections (in the case of the core collections) or core collections (in the case of the mini‐core collections).
Table 1.2. Mini‐core collections formed by Hari Deo Upadhyaya in chickpea, groundnut, pearl millet, pigeonpea, sorghum, and small millet.
Crop
Entire collection
Mini‐core number
% of entire collection
Traits used
Reference
Sorghum
22,473
242
1.08
21
Upadhyaya
et al.
, 2009b
Pearl millet
20,766
238
1.14
18
Upadhyaya
et al.
, 2011 l
Chickpea
16,991
211
1.24
22
Upadhyaya and Ortiz, 2001
Pigeonpea
12,153
146
1.2
34
Upadhyaya
et al.
, 2006e
Groundnut
14,310
184
1.28
34
Upadhyaya
et al.
, 2002a
Finger millet
5,940
80
1.34
20
Upadhyaya
et al.
, 2010c
Foxtail millet
1,474
35
2.37
21
Upadhyaya
et al.
, 2011e
2. Climate‐resilient Germplasm.
Global warming is putting significant stress upon agricultural production and the nutritional quality of staple crops in many parts of the world. Southern Asia and Sub‐Saharan Africa will be the most adversely affected regions, due to climate change and the variability effects. ICRISAT‐mandated crops are widely grown and consumed in these regions (http://faostat.fao.org/site/567/default.aspx#ancor).
The identification and use of climate‐resilient germplasm in crop breeding is the way forward to develop ‘climate‐smart’ crop cultivars. Hari adopted a two‐pronged strategy, first by working with ICRISAT researchers, and second by providing the seeds of several sets of mini‐core collections to NARS partners and working with them to evaluate these subsets for agronomic traits, including stress tolerance. The end result was the identification of several sources of resistance to abiotic and biotic stresses in chickpea and groundnut, with some accessions combining stress resistance and tolerance in good agronomic backgrounds (Upadhyaya et al., 2013a, 2014d). Using a similar approach, Hari and his colleagues identified a number of drought‐tolerant and salinity‐tolerant germplasm accessions in finger millet and/or foxtail millet (Krishnamurthy et al., 2014a, 2014b, 2016).
Blast (Pyricularia grisea) is a devastating disease in pearl millet and finger millet, which has many pathotypes. The work of Hari and his colleagues on screening the pathogenic variability led them to identify accessions that were resistant to multiple pathotypes in pearl millet (Sharma et al., 2015), finger millet (Babu et al., 2013b, 2015) and foxtail millet (Sharma et al., 2014). Downy mildew (Sclerospora graminicola [Sacc.] Schröt) is a highly destructive and widespread disease of pearl millet, while grain mould and downy mildew (Peronosclerospora sorghi) are also important diseases of sorghum. Hari and his colleagues identified a number of accessions with resistance to multiple pathotypes in pearl millet (Sharma et al., 2015) and sorghum (Sharma et al., 2010, 2012). In addition, they identified some lines with good agronomic value, such as early maturity and resistance, and resistance and high seed/fodder yield potential, in both finger millet and pearl millet.
3. Seed Nutrient‐dense Germplasm.
Widespread micronutrient malnutrition in human beings, as a result of deficiency of iron (Fe), zinc (Zn) and β‐carotene, has an enormous socio‐economic cost for society in the developing world (Stein, 2010). Hari saw the need to identify seed nutrient‐dense (i.e. Fe, Zn) germplasm to support crop breeding. After evaluating the mini‐core collections for two seasons, Hari identified a number of different germplasm sources with high seed Fe and/or Zn concentrations in groundnut (Upadhyaya et al., 2012d), pearl millet (Rai et al., 2015), sorghum (Upadhyaya et al., 2016c), finger millet (Upadhyaya et al., 2011d), and foxtail millet (Upadhyaya et al., 2011e). Finger millet and foxtail millet are rich sources of seed protein and calcium (Ca), with some accessions in both of these crops showing exceptionally high protein and Ca contents (Upadhyaya et al., 2011d, 2011e).
4. Bioenergy.
Sorghum is a crop that is used for food, feed, and bioenergy. The stalks are rich in sugar (as measured by Brix). However, the stalk sugar content is greatly influenced by the environment and the crop stage at which the stalks are harvested. Hari evaluated the sorghum mini‐core collection accessions for stalk sugar content for two post‐rainy seasons under irrigated and drought‐stressed conditions. He found that drought stress significantly increased the mean Brix by 12–27%. A few germplasm lines had significantly greater mean Brix (14.0–15.2%), but were agronomically inferior, while some others were agronomically comparable but with similar Brix, such as IS 33844 (Brix, 12.4%) (Upadhyaya et al., 2014a). This indicated that it is possible to select for even higher Brix content in agronomically superior genetic background in germplasm collections. IS 33844 is the local landrace Maldandi that was collected from Maharashtra, India, and it is the most popular sorghum cultivar that is widely grown under decreasing soil moisture conditions during the rabi (post‐rainy) season in India. IS 33844 is tolerant to terminal drought and has excellent grain quality.
5. Germplasm Use in Breeding.
Plant genetic resources are the basic raw materials for genetic progress, and they provide insurance against unforeseen threats to agricultural production. Hari firmly believes that the use of germplasm in crop improvement is one of the most sustainable ways to conserve valuable genetic resources and to broaden the genetic base of crops. Hari partnered with researchers globally to get these subsets (Tables 1–2) evaluated for stress tolerance, yield and seed nutritional traits, and collaborated with molecular biologists to dissect out the population structure and diversity in these representative subsets. This exercise resulted in the identification of several agronomically beneficial and genetically diverse germplasm sources that fulfil the needs of crop breeders. Armed with this valuable information, Hari interacted with crop breeders at ICRISAT and elsewhere, to promote the use of such germplasm in breeding programs.
An analysis of the uptake of germplasm in crop improvement programs at ICRISAT showed that germplasm use has increased since the formation of the mini‐core collections in some crops. For example, there was increased use (≈15% increase) of stress‐tolerant chickpea germplasm during the 2000–2004 and 2005–2009 periods, while in recent years (i.e. 2010–2014), more emphasis (22% increase) has been on the use of germplasm that has agronomic (yield per se) and seed nutritional traits. The trend noted in groundnut was opposite: namely, more emphasis (17% increase) on the use of yield and quality‐enhancing germplasm from 2000–2004, which changed to increased (42% increase) the use of stress‐tolerant germplasm from 2005–2009, with emphasis (46% increase) from 2010–2014 on stress tolerance, yield, and quality enhancement. All of this was possible because of the consistent efforts led by Hari and his colleagues (including those from ICRISAT and NARS countries) to use representative subsets in the identification of new sources of variation with agronomically beneficial traits, and to promote the breeders’ willingness to use new germplasm as a resource in crop breeding.
6. On‐farm Conservation and Use of Diversity.
On‐farm conservation and evaluation of genetic resources provides farmers with the opportunity to select germplasm adapted to their climate conditions. In addition, it also allows evolution of new genetic variants as a result of climate change and variability effects. This facilitates greater and more rapid dissemination of promising seeds among the farming community. Hari’s collaborative work with NARS partners on the evaluation of core/mini‐core collections of finger and foxtail millets, through a project on farmers’ fields in Africa and Asia, provided the farmers with opportunities to access and appreciate the diversity of these neglected crops. Today, farmers own and cultivate some finger millet germplasm sources, such as IE 2440 and 4625 in Uganda, and IE 2872 and 4115 in Kenya, or finger millet (e.g. IE 3575, 4415, 4425, 6045, 6337) and foxtail millet (e.g. ISe 156, 1575) in India. In addition, the NARS partners from these countries have identified stress‐tolerant germplasm that they are using in breeding programs to enhance the genetic potential of these crops.
7. Wild Relatives and Cultigen Genepool.
Wild relatives and their derivatives are sources of variation for agronomic traits, which include stress tolerance, yield, and seed quality. Wild Cicer species, and particularly those from secondary and tertiary genepools that have high levels of resistance to stress tolerance, require vernalization and/or extended day‐length treatments to synchronize their flowering with cultivated chickpea, for interspecific crosses. The use of vernalization and/or photoperiod response enabled Hari and his colleagues to introduce synchronized flowering into a few Cicer species, similar to that of cultivated chickpea (Sharma and Upadhyaya, 2015a). This contributes significantly not only to enhanced use of Cicer species for chickpea improvement, but also to improvements in the regeneration efficiency of Cicer species and their rapid generation turnover.
Cajanus albicans (Wight & Arn.) van der Maesen is a species from the secondary genepool of pigeonpea, and it is known for the long life of its large leaves (leaflet length, 4.4–6.8 cm; leaflet width, 3.1–5.8 cm). Hence, it is an important source of animal feed in semi‐arid tropical regions. It possesses broader pods (9.6–15.0 mm) and high seed numbers (5–7 per pod), is resistant to abiotic (e.g. drought, salinity) and biotic (e.g. pod fly, pod wasp, Alternaria blight, sterility mosaic) stresses, and its high seed protein content (up to 32%) make it particularly attractive (Figure 1.1). Hari had to wait for about 500 days to see the flowering in C. albicans, and another 50–58 days to harvest the mature pods to complete the characterization data on this species. Notably, this produces partial fertile hybrids (Mallikarjuna et al., 2011 and references therein), thus, providing a potential source to broaden the cultigen genepool in pigeonpea.
Figure 1.1.Cajanus albicans, a wild species from a secondary genepool with many desirable characteristics, and a potential source for gene introgression in cultivated pigeonpea.
8. Gaps in Collections.
Identifying gaps in collections and enriching collections with new sources is a critical function of genebank curators. Hari’s work on gap analysis, using geo‐referenced pearl millet landraces from Asian countries (5,768 accessions), revealed parts of the Bihar, Madhya Pradesh, Maharashtra, Rajasthan, and Uttar Pradesh provinces of India as the major geographical gaps in the world collection of pearl millet at ICRISAT (Upadhyaya et al., 2010b).
His similar studies involving pearl millet landraces from southern and eastern Africa (3,750 accessions), and those from west and central Africa (6,434 accessions) also allowed Hari to identify regions in Africa that were not fully represented in ICRISAT collection (i.e. central Sudan and Tanzania, eastern Botswana, west and central Zambia, eastern and central Zimbabwe, southern Mauritania, Niger and Chad and northern Benin, Ghana, and Nigeria) (Upadhyaya et al., 2009c, 2012f). Based on this gap analysis by Hari and requests from NARS partners, the ICRISAT regional genebanks in Africa organized collection missions and collected 6,625 new samples of mandate crops from west and central Africa and southern and eastern Africa regions. These, in my opinion, are important milestones achieved by Hari and his group that further enriched the germplasm collection at ICRISAT.
Hari’s work further revealed that when landraces from the 5°–10°N latitude regions were grown at Patancheru, India, these flowered late and grew tall, and they also produced more tillers. Conversely, those from the 10°–15°N latitude regions had fewer tillers, but with long and thick panicles and larger seeds. Also, landraces from the 10°–15°S and 20°–25°S latitudes are good sources of resistance to bird damage (long‐bristled panicle). Furthermore, Hari found that the landraces of the lower latitude regions (<20°N and S) in both hemispheres are better sources of fodder types (i.e. high tillering, tall, long duration), while those from mid‐latitude regions (15°–20°) in both hemispheres are good sources for enhancing productivity (i.e. early, long and thick panicle, large seeds). Similarly, landraces on both sides of the equator (i.e. within the 10o–20° latitudes) are highly sensitive to the photoperiod (>12.5 hours) and/or temperature (<12 °C), while those from higher latitudes (20°–35°) in both hemispheres showed low sensitivity to both the photoperiod and temperature. The photoperiod and temperature insensitive accessions are represented mostly from the mid‐latitudes (15°–20°) in both hemispheres (Upadhyaya et al., 2012e, 2014f).
B. Molecular Biology and Biometrics
1. Population Structure and Diversity.
Understanding how diversity is structured so as to unlock its potential for crop improvement is an emerging area that has been made possible by rapid advances in the scale, robustness, and reliability of marker technologies, and the sharp fall in the unit costs of their deployment. Hari is probably one of the few CGIAR scientists who used Generation Challenge Program (www.generationcp.org) grants to develop global composite collections, which the molecular biologists at ICRISAT genotyped using high‐throughput assays and simple sequence repeats (SSRs). Hari then used genotyping data and his statistical knowledge to form reference sets in chickpea, pigeonpea, groundnut, pearl millet, sorghum, finger millet and foxtail millet. These reference sets accounted for 78–95% of the allelic variations observed in global composite collections (Table 1.3). Genotyping of reference sets has revealed abundant allelic diversity that grouped the accessions into distinct clusters, with many of the alleles unique in a particular accession in each crop (Upadhyaya et al., 2008a ; Billot et al., 2013). This can be further explored, possibly to associate such allelic diversity with temporal and eco‐geographical diversity, or in proprietary germplasm protection.
Table 1.3. Composite collections and reference sets formed by Hari Deo Upadhyaya in chickpea, groundnut, pearl millet, pigeonpea, sorghum, finger millet, and foxtail millet.
Crop
Number of SSRs used
Composite collection
Reference set
Number of accessions
Number of alleles
Number of accessions
Number of alleles [
n
(%)]
Reference
Sorghum
41
3367
783
383
613 (78.3)
http://genebank.icrisat.org/GB_ReferenceSet/ReferenceSet_Sorghum.aspx
; Billot
et al
., 2013
Pearl millet
19
1021
230
300
218 (94.8)
http://genebank.icrisat.org/GB_ReferenceSet/ReferenceSet_Pearlmillet.aspx
Chickpea
48
2915
1683
300
1315 (78.1)
http://genebank.icrisat.org/GB_ReferenceSet/ReferenceSet_Chickpea.aspx
; Upadhyaya
et al
., 2008b
Pigeonpea
20
952
197
300
187 (94.9)
http://genebank.icrisat.org/GB_ReferenceSet/ReferenceSet_Pigeonpea.aspx
Groundnut
21
852
490
300
466 (95.1)
http://genebank.icrisat.org/GB_ReferenceSet/ReferenceSet_Groundnut.aspx
Finger millet
20
959
231
300
206 (89.2)
http://genebank.icrisat.org/GB_ReferenceSet/ReferenceSet_Fingermillet.aspx
Foxtail millet
19
452
362
200
316 (87.3)
http://genebank.icrisat.org/GB_ReferenceSet/ReferenceSet_Foxtailmillet.aspx
2. Genome‐wide Association Mapping.
The diversity panels of germplasm collections, such as the conventional core and mini‐core collections, or genotype‐based reference sets, are ideal germplasm resources for studying linkage disequilibrium and association mapping in crop plants. Identification of candidate genes associated with abiotic stress responses will accelerate breeding efforts that are aimed at enhancing productivity in drought‐stressed environments.
Hari’s collaborative work with molecular biologists led to the identification of 18 single nucleotide polymorphisms (SNPs) in chickpea reference accessions that were significantly associated with drought‐avoidance root traits, carbon isotope discrimination, heat tolerance, harvest index, and 100‐seed weight under drought‐stressed conditions (Roorkiwal et al., 2014a). Similar work using SSRs in groundnut reference accessions revealed significant marker‐trait associations for drought‐tolerance traits (e.g. chlorophyll readings, harvest index) and seed weight, under both well‐watered and drought‐stressed conditions (Pandey et al., 2014).
Together with his US collaborators, Hari identified significant marker‐trait associations for phenology, panicle architecture and stress tolerance in sorghum, using diversity panel germplasm (Lasky et al., 2015 ; Morris et al., 2013 ; Upadhyaya et al., 2012 g, 2012 h; Upadhyaya et al., 2013f, 2013 g, 2013 h; Upadhyaya et al., 2016f), which also included the sorghum mini‐core collection accessions formed at ICRISAT (Upadhyaya et al., 2009b).
3. Candidate Genes Associated with Agronomically Useful Traits.
The discovery of large‐scale high‐quality SNPs led Hari and his collaborators to identify candidate genes that can regulate complex agronomic traits, such as flowering (e.g. efl1, FLD, GI, Myb, SFH3, bZIP, bHLH, SBP) (Upadhyaya et al., 2015a), plant height (Kujur et al., 2016), number of branches (e.g. PIN1, TB1, BA1/LAX1, GRAS8, ERF, MAX2, lipase) (Bajaj et al., 2016b), number of pods and seeds per plant (Kujur et al., 2015b), seed colour (e.g. MATE) (Bajaj et al., 2015a), 100‐seed weight (e.g. CSN8, ERF, TF) (Bajaj et al., 2015b ; 2015c, 2016a ; Das et al., 2015a), and seed protein (e.g. zinc finger transcription factor) (Upadhyaya et al., 2016a) in chickpea.
4. Ethnolinguistic Groups Shaped Sorghum Diversity in Africa.
Sub‐Saharan Africa, and more particularly Ethiopia, Sudan, and Chad, were the primary centres of the origin and domestication of sorghum. Hari’s collaboration with researchers from Norway and Sudan highlighted three major sorghum populations that were associated with the distribution of ethnolinguistic groups in Africa (Westengen et al., 2014). The co‐distribution of the central sorghum population and the Nilo‐Saharan language family demonstrated a close and causal relationship between the distribution of sorghum and languages in the region between the Chari and Nile rivers. The southern sorghum population was associated with the Bantu languages of the Niger‐Congo language family, while the northern sorghum population was distributed across the early Niger‐Congo and Afro‐Asiatic language family areas with dry agroclimatic conditions. Furthermore, the genetic structure within the central sorghum population was associated with language group expansions within the Nilo‐Saharan language family.
The Western‐Nilotic ethnolinguistic group (e.g. the Pari people) provided a window into the social and cultural factors involved in the generation and maintenance of the continent‐wide diversity patterns. The age‐grade system is a cultural institution that was important for the expansive success of this ethnolinguistic group in the past, and it had a central role in the management of sorghum landraces. This continues to underpin the resilience of their traditional seed system, which supports the ‘farming‐language co‐dispersal hypothesis’, which proposes that farming and language families have moved together through population growth and migration (Diamond and Bellwood, 2003 ; Jobling et al., 2013).
5. Genome Sequencing.
Hari was involved in the sequencing of the reference genomes of chickpea (Varshney et al., 2012b), pigeonpea (Varshney et al., 2013), groundnut (Chen et al., 2016) and pearl millet. Hari’s interest in genome sequencing lies in finding and associating sequence variations that have agronomically beneficial traits. He believes these will improve the efficiency of genebank operations, with particular reference to conservation, regeneration and use of germplasm with unique characteristics in crop breeding. Hari is participating in the 3000‐genomes project of chickpea, using a composite collection and 300‐groundnut genomes with reference set accessions. He hopes that this will provide him with many opportunities to use sequence variations for efficient management and use of genetic resources.
C. Groundnut Breeding
At ICRISAT Patancheru India, Hari was initially involved in developing early‐maturing (short duration; 90 days in the rainy season) and aflatoxin‐resistant groundnut varieties. For a few years (2000–2001 to 2004–2005), Hari also managed the groundnut breeding research at ICRISAT Regional Research Centre, Lilongwe, Malawi. His focus in Malawi was to incorporate early‐maturity and resistance to leaf spots and rosette disease into locally adapted cultivars from the eastern and southern African areas. In 2012, Shyam N. Nigam, Principal Scientist and leader of the Groundnut Breeding Unit, ICRISAT, retired from ICRISAT, and the management asked Hari to shoulder additional responsibility as Principal Groundnut Breeder to mentor the incumbent and to continue working on breeding for early‐maturity, drought tolerance, aflatoxin resistance, and high oil content in groundnut.
1. Early Maturity.
Developing early‐maturing (i.e. 90 days), high‐yielding groundnut varieties was one of several breeding objectives when groundnut was included in ICRISAT as one of its mandate crops in 1976. In the early days, the often‐used early maturing source was a small‐seeded germplasm line, Chico (ICG# 476), from Russia (Bailey and Hammons, 1975), although it provided only limited success in breeding early maturity into an improved genetic background. The most common breeding strategy adopted during that time was to plant segregating populations and harvest them when the crop received 1470° Cd (cumulative thermal time), which was equivalent to 90 days after sowing during the rainy season in Patancheru, India, while the advanced lines were stagger‐harvested when the crop accumulated 1240° Cd (equivalent to 75 days during the rainy season) and 1470° Cd after sowing in Patancheru, India (Vasudeva Rao et al., 1992).
The three things Hari did differently from his predecessors were:
he diversified the sources of early maturity by identifying new and diverse ones (Upadhyaya
et al
., 2006f);
he investigated the genetics of early flowering (first flower appearance, accumulation of 25 flowers) (Upadhyaya and Nigam, 1994); and
he started selecting for early maturity on an individual plant basis, instead of adopting bulk selection method of breeding, as was the practice during these early days.
Hari used segregating populations, harvested at pre‐designated heat units (1470 °Cd) in Patancheru, India. He then imposed other selection criterion: selection based on percentage sound matured kernels (SMK%) and on uniformity of seed size and shape (Legumes program Annual Report, 1993). This two‐way selection pressure paid rich dividends and, in a short time, he developed a large number of early maturing advanced varieties with potential yield as high as 3 tons ha–1 to 4 tons ha–1, harvested at 1470 °Cd in Patancheru, India (ICRISAT Archival Report of Research Program on Grain Legumes 2012–2013, 2014).
Selecting for high oil content or large‐seed size in an early maturity background with improved yield potential was a breeding challenge. Hari was, however, successful in combining early maturity and high oil content, or early maturity and large‐seed size, in improved genetic backgrounds (ICRISAT 2014, 2015 ; Upadhyaya et al., 2005b).
Ultra‐susceptibility to foliar diseases (i.e. rust, late leaf spot) and lack of fresh seed dormancy were associated with early maturity, with the potential risk to seed germination if it rained during the harvest and if there was a delay in lifting the crop from the field. Hari was successful in combining early maturity and fresh seed dormancy (Upadhyaya et al., 1997a, 2001e), or tolerance to rust and late leaf spot (Upadhyaya et al., 2001d) in some varieties.
Post‐rainy season groundnut in India coincides with early winter in peninsular India, and there was a need to combine cold tolerance at germination in an early maturing genetic background. Here, Hari used his remarkable good sense and sharp mind. On one fine morning during the winter season, he visited his groundnut breeding field together with his technician, as the crop was just emerging from the ground. He noted that some plots had early emergence while, in many others, even the soil crust was not broken.
Together with his technician, he noted these plots, and he regularly visited them for about two weeks. He observed that some lines also had greater seedling vigour in addition to early emergence, compared with the others. Both early emergence and greater seedling vigour helped Hari to identify cold‐tolerant lines in groundnut. Following this, he picked up a breeding line, ICGV 92267, that combined early maturity, low temperature tolerance at germination (≰12 °C), and resistance to rust and late leaf spot (Upadhyaya et al., 2002c). For many aspects, I consider that this was a remarkable development, to enhance the adaptation of groundnut in new areas where the temperature is low at planting.
2. Drought Tolerance.
Hari carried forward the drought tolerant breeding populations/advanced breeding lines developed by Shyam N. Nigam to their logical conclusion, while also introducing new elements to enhance the drought tolerance in groundnut. For example, he identified additional sources of drought tolerance in good agronomic backgrounds (ICRISAT 2014, 2015), which he crossed with early maturing varieties to produce new breeding populations that combined early maturity with drought tolerance.
Hari continued to evaluate breeding populations/advanced varieties under rain‐fed (i.e. no supplemental irrigation) and irrigated (i.e. fully irrigated) conditions in rainy seasons (June/July to October/November), and under irrigated and stress conditions (i.e. withholding alternate irrigations from 60 days after sowing) in post‐rainy (October/November to March/April) seasons. He also selected varieties that showed high pod yield under drought‐stressed environments, with no yield penalty under favourable environments. Hari developed a number of high‐yielding and drought‐tolerant varieties (ICRISAT 2014, 2015), and some of these are already in evaluation trials in national programs across Asia and Africa.
3. Aflatoxin Resistance.
Aflatoxin contamination is a serious quality problem in groundnut, and this involves pod, seed coat and cotyledons as the three components:
resistance to natural seed infection;
in vitro
seed colonization; and
aflatoxin production by
Aspergillus flavus
needs to be combined (Nigam
et al
., 2009).
The environmental factors (e.g. drought stress at time of pod development and maturity) and post‐harvest processing (i.e. harvesting, drying, curing) and storage (i.e. mainly temperature, humidity) conditions largely influence the aflatoxin contamination in groundnut. Hari was successful with the combination of resistance to natural seed infection and/or in vitro seed colonization by A. flavus into an improved genetic background, some of which supported very low levels of aflatoxin contamination, with potential yields of 2.5 t ha–1 to 4.0 t ha–1 during the rainy and post‐rainy seasons in Patancheru, India. However, he agreed that much more needs to be done (ICRISAT 2014, 2015 ; Rao et al., 1995 ; Upadhyaya et al., 2001c).
4. Farmers Participatory Varietal Selection.
Hari firmly believes that the success of plant breeders depends on how they perceive what farmers need and how close the breeders can integrate such traits into new varieties. The opportunity to work with stakeholders (i.e. the farmers, consumers, traders) adds additional value to successful plant breeders. The Tropical Grain Legume project II (Monyo and Gowda, 2014), which was supported by the Bill and Melinda Gates Foundation (www.gatesfoundation.org), provided such a platform to many scientists in the CGIAR and NARS institutions, to test products/technologies under on‐farm conditions with active participation from the farmers.
Working with farmers in India, Hari noted they preferred varieties that mature early, are stress‐tolerant, have ease of harvest and shelling, and have uniformity of pod and seed characteristics. Three of the groundnut varieties among those bred by Hari were selected by the farmers, and later these were released in India: ICGV 00350 in Andhra Pradesh and Tamil Nadu; ICGV 00351 as ‘CO 7’ in Tamil Nadu; and ICGV 93468 as ‘Avatar’ in Uttar Pradesh. The first two were selected for drought tolerance and adaptation to both rain‐fed and irrigated post‐rainy season production systems, while the last one was adapted to spring season cultivation (February/March to June/July), especially after a harvest of potato in Uttar Pradesh.
This ‘Avtar’ cultivar has revolutionized groundnut production during the spring season in the Uttar Pradesh state of India. The acreage has expanded from a mere 20,000 ha in 2004 to 317,068 ha in 2011, and production during the same period increased from 53,100 t to 802,616 t; however, the average productivity remained at 2.53 t ha–1. Shyam N. Nigam of ICRISAT and officials from the Department of Agriculture, Uttar Pradesh, India, contributed to the popularizing of ‘Avtar’ for cultivation during the spring season in Uttar Pradesh. Hari is now using the knowledge gained from the interaction with the farmers in the development of new cultivars with the traits preferred by the farmers, and he has several such products (varieties) in his cupboard that are waiting for the opportunity for on‐farm evaluation and selection by the farming community.
D. Chickpea Breeding
Wilt caused by Fusarium oxysporum f.sp. ciceris is widespread, and causes substantial yield loss to chickpea production worldwide (Nene et al., 1978). Early studies on the pathotypes revealed four races of wilt (Haware and Nene, 1982), with resistance to race 1 prevalent at Patancheru, which is controlled by a single recessive gene in crosses involving C 104 as the susceptible parent (Kumar and Haware, 1982). However, in the crosses with cultivar JG 62 as a susceptible parent, the number of susceptible plants was too large to give a good fit to the expected ratio.
Hari’s seminal work on inheritance of wilt resistance revealed that the appearance of the wilting symptoms differed amongst the JG 62 and C 104 susceptible parents. C 104 wilts later than JG 62, whereby the difference in the time of wilting is controlled by a single gene, with early wilting partially dominant to late wilting (Upadhyaya et al., 1983a). Here, further studies by Hari showed that the resistance is controlled by at least two genes, both of which must be present in their homozygous recessive forms to impart complete resistance (Upadhyaya et al., 1983b) although, individually, the genes delay wilting, as in C 104. This discovery by Hari laid a strong foundation later on for the wilt resistance breeding program, which was one of the most successful chickpea disease‐resistance breeding programs at ICRISAT, and indeed, elsewhere.
