Plant Breeding - 2024 - Nagaraja - Panorama of Small Millets Breeding A Review

Received: 27 January 2024 Revised: 6 June 2024 Accepted: 9 July 2024
DOI: 10.1111/pbr.13211
REVIEW ARTICLE
Panorama of small millets breeding: A review
T. E. Nagaraja 1 | S. Gazala Parveen 1 | J. Meenakshi 1 | S. Murtujasab 1 |

Chetana 1 | M. Manjunatha 1 | Sujata Bhat 1 | C. Nandini 2
1
ICAR-All India Coordinated Research Project
(Small Millets), Project Coordinating (PC) Unit, Abstract
University of Agricultural Sciences, Bangalore,
Small millets, also known as nutri-cereals, belong to the grass family Poaceae that
India
2
Zonal Agricultural and Horticultural Research have tiny seeds and are drought and heat tolerant, besides being powerhouse of
Station (ZAHRS), Babbur Farm, Hiriyur, Keladi nutrition, hence making them ideal crops that shall combat not just food and nutri-
Shivappa Nayaka, University of Agricultural
and Horticultural Sciences, Shivamogga, India tional security but also adapt well to the changing climatic conditions. They grow well
on marginal soil and require least resources for their cultivation. Nutritional superior-
Correspondence
T. E. Nagaraja, ICAR-All India Coordinated
ity to popularly cultivated major cereals makes them a magnificent alternative food
Research Project (Small Millets), Project especially in the age of lifestyle related disorders. In this review, we provide insights
Coordinating (PC) Unit, GKVK Campus,
University of Agricultural Sciences, Bangalore,
of the initial commencement of research on small millets and deliberate the breeding
Karnataka 560 065, India. progress based on conventional and genomics-based approaches. The small millets
Email: [email protected]
under discussion are finger millet, foxtail millet, proso millet, little millet, barnyard mil-
let, kodo millet and browntop millet. The major focus here is to recapitulate the
breeding activities in the above-mentioned millets rather than providing a way for-
ward for breeding small millets.
KEYWORDS
breeding, crop improvement, hybridization, omics, small millets
1 | I N T RO DU CT I O N minor millets/small millets (Gupta, 2006).Millets thrive well in low

rainfall areas with marginal fertile soils, adapt to varied ecological
The grass family Poaceae includes cereal food crops with small grains conditions, require minimum inputs for their growth, incidence of
known as millets (Marcon, 1994). They form an agronomic group insect pests and diseases is low and are considerably less prone to
rather than a taxonomic group. Owing to their nutritional profile that environmental burdens, hence they are also regarded as climate-
is able to meet the requisites for regular functioning of human body, resilient crops (Bandyopadhyay et al., 2017; Goron & Raizada, 2015;
widely, they are also known as ‘nutri-cereals’. The abundance of Saxena et al., 2018). About 80% of millets produced are utilized as
both wild and cultivated forms of millets especially in tropical west- food by humans, followed by as poultry feed, potable alcohol and for
ern Africa as compared with many other parts of the world makes it industrial purpose. Their budding crucial use as an industrial raw
their evolutionary centre of origin. Based on the per cent area grown material in making biscuits, confectioneries, beverages, weaning
and the size of the grain, they are categorized as major millets and foods and beer (Anukam & Reid, 2009; Laminu et al., 2011) is gaining
minor millets. Sorghum (Sorghum bicolor [L.]) and pearl millet (Pennise- significance. Despite millets being primeval food crops to ancient
tum glaucum [L.] R. Br.) are major millets, while finger millet (Eleusine people, the green revolution ramifications resulted in their decreased
coracana [L.] Gaertn.), foxtail millet (Setaria italica [L.] P. Beauv.), little cultivation, thereby impeding their usage and importance in human
millet (Panicum sumatrense Roth. ex. Roem. & Schult.), proso millet diet. Lately, the aftermath of current human food habits and the
(Panicum miliaceum L.), kodo millet (Paspalum scrobiculatum L.), barn- resultant lifestyle has resulted in burgeoning human diseases like dia-
yard millet (Echinochloa crusgalli [L.] P. Beauv. and Echinochloa colona betes, hypertension and cardiovascular issues; consequently, urging
[L.] Link) and brown top millet (Brachiaria ramosa [L.] Stapf. sustainable and viable diet options with the inclusion of millets is
= Urochloa ramosa [L.] T.Q. Nguyen) (Table 1) are designated as highly encouraged.
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2 NAGARAJA ET AL.
TABLE 1 Details of small millets discussed in this review.
Chromosome
number and Genome Major areas of
Crop Origin Progenitor ploidy size (in Mb) production Reference
Finger millet (Eleusine East Eleusine 2n = 4x = 36 1196 (ML India, Ethiopia, Nepal, de wet et al. (1984);
coracana [L.] Gaertn.) African indica Eleusine 365) and Uganda, Malawi, Hilu et al. (1979); Hilu
highlands floccifolia/Eleusine 1500 (PR Burundi, Sri Lanka and and de Wet (1976);
tristachya 202) Rwanda Hittalmani et al. (2017);
Hatakeyama et al.
(2018)
Foxtail millet (Setaria China Setaria viridis 2n = 2x = 18 510 (Yugu 1) China, Myanmar, India Li et al. (1945); Yang
italica [L.] P. Beauv.) and 423 and Eastern Europe et al. (2012); Wang
(Zhang gu) et al. (1995); Diao and
Jia (2017)
Little millet (Panicum India Panicum psilopidium 2n = 4x = 36 139,384 bp India de Wet et al. (1983);
sumatrense Roth.) (0.139 MB) Sebastin et al. (2018)
Proso millet (Panicum China Panicum capillare 2n = 4x = 36 923 Russia, United States of de Wet (1986); Hunt
miliaceum L.) and Panicum repens America (USA), South et al. (2011); Hunt et al.
Korea, Kazakhastan, (2008); Lu et al. (2009);
France, Poland, Belarus, Zou et al. (2019)
India and Iran
Kodo millet India Paspalum sanguinale 2n = 4x = 40 - India Christopher et al.
(Paspalum (1987); de Wet
scrobiculatum L.) (1983a); Hiremath and
Dandin (1995)
Barnyard millet Peninsular Echinochloa 2n = 6x = 54 1270 India, Japan, China and de Wet (1983); Guo
(Echinochloa crusgalli India crusgalli/Echinochloa Malaysia et al. (2017)
[L.] P. Beauv. and colona
Echinochloa esculenta
[A. braun] [L.] Link)
Browntop millet South - - - - Clayton et al. (2006);
(Brachiaria ramosa [L.] India/ Kingwell-Banham and
Stapf. = Urochloa South Fuller (2014)
ramosa [L.]) India
Note: -: Data are not available.
By and large, millets are storehouse of fibre, minerals and micro- poly-unsaturated fatty acids and flavonoids including magnesium,
nutrients (Pramitha Lydia et al., 2021). The nutritional constitution of phosphorous and protein content in little millet is noticed (Indirani &
small millets is detailed in Table 2. Millets differ extensively in constit- Devasena, 2021), while essential amino acids like lysine, leucine, iso-
uents like carbohydrates, as proportion of amylose and amylopectin leucine and methionine are more common in proso millet (Pramitha
ranges from 16% to 28% and 72% to 84%, respectively. They also Lydia et al., 2021). Increased phenol content in kodo millet aids in rad-
possess abundant vitamins, namely, thiamine, riboflavin, folic acid and ical scavenging activity (Deepak Taggelli & Thakur, 2018). Highest die-
niacin. They have high anti-oxidants and phytochemicals that protect tary fibre and niacin content among all the small millets is present in
against the perils of heart disease and diabetes, aid in easy digestion, barnyard millet; it also has high phosphorous and magnesium profile
lower the threat of cancer, detoxify the body, increase immunity, (Renganathan et al., 2019). Browntop millet is rich in phytochemicals
improve muscular and neural systems and provide defence against such as flavonoids, quinones, tannins and resin (Singh et al., 2022).
variety of degenerative diseases like metabolic syndrome and Parkin- Millets are comparable with rice and wheat in terms of nutritional pro-
son's disease (Chandrasekara & Shahidi, 2012; Manach et al., 2005; file, and rich in some of the minerals and fatty acids, that makes them
Rao et al., 2011; Scalbert et al., 2005). Their non-glutinous nature a suitable substitute of rice and wheat.
makes them a safe food choice for people experiencing gluten allergy Humans have been directly utilizing nearly 80,000 plant species
and celiac disease. Higher levels of calcium, phosphorous, iron and for food, fodder, fibre, medicinal and industrial purposes, among
zinc content in finger millet, additionally a higher caloric value due to which 25,000 are edible (Muthamilarasan et al., 2019). Currently,
its carbohydrate content, were reported (Pore & Magar, 1979). Proso mere 30 plant species are in cultivation that meet the food require-
millet has a similar protein composition (11.60% of dry matter) to ment of eight billion people (https://www.fao.org/faostat/en/#data/
wheat, but it has more of the essential amino acids (leucine, isoleu- OA); despite this, nearly 10% of the world's population remains
cine, and methionine) (Kalinova & Moudry, 2006). The presence of malnourished (https://www.statista.com/statistics/264900/number-
NAGARAJA ET AL. 3
TABLE 2 Nutrient composition of small millets (per 100 g).
Component Finger millet Foxtail millet Little millet Proso millet Kodo millet Barnyard millet Browntop millet
Moisture (g) 10.87 ± 0.61 9.59 ± 0.61 14.23 ± 0.45 7.25 ± 0.37 14.23 ± 0.43 - 11.90
Protein (g/100 g) 7.70 12.30 8.92 ± 1.09 12.50 8.92 ± 1.09 6.20 8.89
Ash (g) 2.04 ± 0.34 3.30 1.72 ± 0.27 1.90 1.72 ± 0.27 2.20 -
Total fat (g/100 g) 1.02 ± 0.14 4.50 2.55 ± 0.13 1.10 2.55 ± 0.13 2.20 1.89
Crude fibre (g) 3.60 8.00 6.39 ± 0.60 5.20 6.39 ± 0.60 13.60 8.20
Carbohydrate (g) 66.82 ± 0.73 71.29 65.55 ± 1.29 70.04 66.19 ± 1.19 65.55 71.32
Energy (kcal) 328 331.00 341.00 341.00 309.00 307.00 338.00
Thiamine B1 (mg/g) 0.37 ± 0.041 0.59 0.24 ± 0.042 0.41 0.29 ± 0.054 0.33 -
Chromium (mg) 0.032 ± 0.019 0.03 0.016 ± 0.006 0.02 0.021 ± 0.027 0.09 -
Copper (mg) 0.67 ± 0.22 1.40 0.34 ± 0.08 1.60 0.26 ± 0.05 0.60 1.23
Riboflavin B2 (mg) 0.17 ± 0.008 0.11 0.05 ± 0.008 0.28 0.20 ± 0.018 0.28 -
Niacin B3 (mg) 1.34 ± 0.0021 3.20 1.29 ± 0.02 4.20 1.49 ± 0.08 4.20 -
Calcium (mg) 344.00 36.00 16.10 14.00 15.30 20.00 28.00
Phosphorus (mg) 250.00 290.00 220.00 206.00 188.00 280.00 276.00
Mg (mg) 130.00 81.00 133.00 81.00 122.00 137.00 94.50
Zn (mg) 2.50 2.40 3.70 2.36 1.60 3.00 2.75
Fe (mg) 4.60 2.80 1.20 0.80 0.80 5.00 7.72
Note: -: Data are not available.

Source: Indian Food Composition Tables, NIN-2007; Nutritive Value of Indian Foods; Roopa (2015); Shankaramurthy and Somannavar (2019).
of-undernourished-starving-people-worldwide/). With this downfall, minimal dependency on synthetic fertilizers and also carbon footprint
the aim to achieve the Sustainable Development Goal 2 (‘Zero Hun- of millets is relatively lower than major cereals (Saleh et al., 2013).
ger’) by 2030 seems uncertainS. Additionally, owing to rise in global Therefore, promotion of millet cultivation by way of developing high-
mean surface temperature, remarkable yield loss in many commercial yielding cultivars that are not just resistant to abiotic and biotic stres-
crops is predicted (Avnery et al., 2011; Challinor et al., 2014; Teixeira ses but are also less inclined to lodging and shattering is the pressing
et al., 2013), consequently leading to heat and drought stresses and priority, as majority of the currently grown cultivars are selections
related abiotic and biotic stresses. Furthermore, turmoil in global sup- from available germplasm accessions that were bred decades ago and
ply chain, restrictions on the export of agricultural commodities have reached a yield plateau. In this article, we aim to review the pro-
accompanied with inflation and various economic consequences as a gress made in breeding small millets using conventional and
result of the aftermath of the pandemic COVID-19 demands an action genomics-based approaches thus far.
plan to address food insecurity (Mehanathan & Manoj, 2021) and cor-
roborate preparedness for such catastrophes by regulating the pro-
duction of nutrient dense crops that give higher economic yield and 2 | S T R A T EG I E S T O B R E ED SM A LL
are less resource demanding even in harsh climatic conditions. Given MILLETS
these umpteen challenges, an arduous task awaits the plant breeders
to develop crop varieties that not just meet the nutritional require- For the longest time since the cultivation of millets, morphological
ments of humans but also keeps in check the global mean surface selection of superior plants from local landraces or germplasm acces-
temperature. Millets being highly nutritious crops that require very lit- sions were utilized to grow in the subsequent seasons. It is only after
tle inputs for their cultivation shall serve as suitable alternative to the the establishment of various research centres dedicated exclusively to
currently grown rice, wheat, maize, potato, soybean and sugarcane breed improved small millets cultivars that the initiatives to identify
that share more than 75% of total plant-derived energy intake. There- and develop better cultivars with superior yield levels were attempted
fore, the above-mentioned scenarios encourage the cultivation and using conventional breeding methods. Considering the floral biology
consumption of small millets, because they are preferable to major of a crop, duration of flowering period and pollination behaviour are
cereals in terms of agroecological traits (Fuller et al., 2004) and nutri- of paramount importance to design a suitable and efficient breeding
tional quality and possess the potential to have solution for food secu- programme aimed to improve yield stability and productivity. Small
rity. Additionally, millet cultivation should be promoted as it ensures millets are highly self-pollinated; the floret size in small millets is tiny,
stable income to growers in case of the failure of main crop, their cul- impeding the task of emasculation to attempt hybridization (Gupta
tivation encourages sustainable agricultural practices as they have et al., 2011; Nagaraja et al., 2023). Taxonomically, the floral biology
4 NAGARAJA ET AL.
and the process of flowering is incomprehensible. As a result, majorly, subsequent seasons in Africa (Jameson, 1970). To encourage finger
the breeding methods routinely employed in self-pollinating crops can millet research, Serere Experimental Station (SES), which was later on
be employed in small millets as well like pure line selection, mass renamed as Serere Research Station (SRS) in 1970, was established
selection, pedigree selection, bulk breeding method, single-seed under the umbrella of East African Agriculture and Forestry Research
descent method, backcross breeding, heterosis breeding and mutation Organization in 1965 (East African Agriculture and Forestry
breeding. Exploitation of hybrid vigour in small millets is limited owing Research Organization, 1971). Initially, the focus of SES was largely
to the obstacles in hybridization. Comprehending the floral biology on germplasm collection, cleaning and safe-guarding the farmer-
and pollination system aids to generate appropriate emasculation and preferred landraces. Cytoplasmic male-sterile (CMS) lines from Inter-
crossing protocols. Therefore, attempts to develop crossing tech- national Crop Research Institute on Semi-Arid Tropics (ICRISAT) were
niques that shall result in successful rate of hybridization is highly introduced in Uganda in 1970's to use as parents in hybridization pro-
recommended (Nagaraja et al., 2023). Additionally, generation of grammes. This resulted in recognition of a male-sterile line Gulu
male-sterile lines would be a practicable choice to exploit heterosis. E. Simultaneously, crossing, selfing and double crosses were
Routinely, in self-pollinated crops, mutation breeding has played an attempted, wherein backcross programme largely focused on blast
indispensable role where hybridization is difficult to establish variabil- and lodging resistance (East African Agriculture and Forestry Research
ity. Also, the use of chemical hybridizing agents (CHAs) to effectuate Organization, 1971). In 1969–1971, three varieties, namely, Engeny,
male sterility in small millets needs to be investigated. The adequacy Gulu E and Serere 1, which had farmer-preferred traits like good malt-
of mutagens relies on its mutagenic effectiveness (mutations per unit ing and grain quality, lodging resistance, good yield level and were
dose of mutagen) and efficiency (mutation in relation to undesirable adaptable to different environments, were delivered for commercial
changes/damage like sterility, lethality and injury) to generate wide cultivation (Esele, 1989).
range of numerous and desirable mutants. This was followed by the release of Pese 1 (P224), Seremi 1
(P249), U15 (Seremi 2), Seremi 3 (S x17-88), Serere 14 and Sec 695 in
the 1980s (Esele, 1989). The insurgence (1986–1992) in eastern
2.1 | Throwback to initiation of small millets Uganda led to mass exodus of scientists from SRS to Namulonge
research Research Station (NRS) and financial constraints for smooth conduct
of research (Doggett, 1989), resulting in crumpling of finger millet
Conventional breeding methods like pureline selection and pedigree breeding. Thereafter, SRS was renamed Serere Agricultural and Ani-
breeding for varietal release in majorly cultivated crops except small mal Research Institute (SAARI) with the mandate to conduct finger
millets were prominent since the 1900s; it is only in the early 1950s millet breeding under the cereals improvement programme
that improvement in small millets gained recognition (Paroda & (Oryokot, 2001). In 2017, five finger millet varieties, namely, Naromil
Mal, 1989). Since, self-pollination is a rule in small millets; often, 1 (FMS383), Naromil 2 (FMS 376), Naromil 3 (FMS 72),
locally available accessions were evaluated for line development by Naromil 4 (SEC 915) and Naromil 5 (IE 2440), were released for com-
plant breeders. Progressively, besides pureline selection and pedigree mercial cultivation (Kasule et al., 2023).
breeding, mass selection and mutation breeding were also attempted Breeding activities in foxtail millet commenced with correlation of
in small millets. Until the 1950s, the research centres across India grain yield among landraces in the 1950s and 1960s; eventually,
relied on the utilization of locally adapted finger millet lines for its breeding methods like single-ear and bulk selection were attempted.
genetic improvement. Subsequently, after the establishment of Pro- Similar to finger millet, morphologically superior foxtail millet lines
ject for Implementation of Regional Research on Cotton, Oilseeds and were encouraged for commercial cultivation after evaluation at field
Millets (PIRRCOM) in the late 1950s which also lead to the function- level. In India, it was in 1940 that the Tamil Nadu Agricultural Univer-
ing of All India Coordinated Millets Improvement Project (AICMIP) in sity (TNAU) initiated foxtail millet breeding, followed by Agricultural
1965, that undertakes major efforts towards improvement in small Research Centre (ARS) (Hagari) in Karnataka that bred H-1 and H-2,
millets. The AICMIP functions in collaboration with various voluntary which were tall but prone to lodging with lower yield levels. Till date,
centres located across Indian State Agricultural Universities like Ber- pureline selection and hybridization followed by selection have been
hampur (Orissa), Coimbatore (Tamil Nadu), Dindori (Madhya Pradesh), followed to breed 40 improved foxtail millet varieties in India. Like-
Jagdalpur (Chhattisgarh), Kolhapur (Maharashtra), Ranchi (Jharkhand) wise, in 1940, the proso millet improvement programme commenced
and Rewa (Madhya Pradesh), while the lead centre is situated in Uni- in China in Agricultural Experiment Farms of the former province of
versity of Agricultural Sciences, Bangalore, India, whose mandate is to Suyuan (currently Langshan town of Linhe District, Inner Mongolia)
conserve, evaluate, characterize, document, identify and supply and the Guanghua Farm (the predecessor of the Yan'an Research
aspects of small millets. Institute of Agricultural Sciences) of the Shan-Gan-Ning Region, while
The second largest finger millet growing area in the world after the Rajendra Prasad Agricultural University in Bihar, India, initiated
Karnataka, India, is Eastern and Southern Africa particularly in Kenya, proso millet research in 1960. Lenis Nelson, in 1972, started exclusive
west of Rift Valley and Lake Victoria region (National Research breeding programme for proso millet productivity improvement under
Council, 1996). Selection and maintenance of superior fingers the alternative crops breeding programme at Panhandle Research and
obtained from indigenous cultivars formed the basis of seed for Extension Center (PHREC) in the United States. Consequently,
NAGARAJA ET AL. 5
considerable improvement in proso millet productivity was evident In 1931, the College of Agriculture, Hebbal campus, Bangalore,
with the release of four cultivars, namely, Sunup, Dawn, Cerise and initiated finger millet improvement programme, which eventually
Rise. The major breeding methods employed in proso millet breeding developed and released the varieties PR-202 and HR-911. Later on,
are line selection and cross-based pedigree selection. Very meagre to the AICRP (Small millets) at University of Agricultural Sciences Banga-
no information is available on breeding methods adapted to develop lore, Karnataka, was established with the mandate to develop high-
little millet cultivars. Because the floral biology of proso millet and lit- yielding and nutrient-rich small millet varieties suitable to both rainfed
tle millet are similar, the strategies followed to breed the respective and irrigated ecosystem. As a result, several varieties, namely, GPU-
cultivars is also similar. The small floret size of barnyard millet ham- 26, GPU-28, GPU-45, GPU-48, GPU-66 and ML-365, were identified
pers the hybridization programme, making it more tedious and released for commercial cultivation. The recent and popular small
(Gupta, 2006; Nagaraja et al., 2023). As a result, barnyard millet millet varieties bred by AICRP (Small millets) are presented in Table 3.
breeding programmes largely rely on traditional mass selection and
pureline selection. Until now, 23 barnyard millet varieties have been
released following pureline selection and hybridization followed by 2.2 | Conventional breeding methods employed in
selection in India. The TNAU initiated the research on kodo millet in small millets
1940s, subsequently developing PLR-1, that was a low yielder and
late in maturity. Till this date, pureline selection and mutation breed- 2.2.1 | Introduction
ing were followed to release 45 kodo millet varieties in India. Negligi-
ble breeding progress is witnessed in browntop millet, the only Reports on introductions in small millets are not available except fox-
commercially available browntop millet cultivars in India are HBr-2, tail millet. In Korea, the varieties Yonyi and Jigu No. 2 were selected
rich in iron (70 mg/kg), zinc (42 mg/kg) and crude fibre (13%) devel- for direct use.
oped as selection from germplasm accession and GPUBT-2
(Figure 1a,b) resistant to leaf blight disease developed from AICRP
(Small millets), University of Agricultural Sciences, Bangalore. In India, 2.2.2 | Mass selection
of the 323 varieties of seven small millets (Figure 2), about 65% are
selections from landraces, 30% are pedigree selections and 5% The seeds from morphologically appealing plants in a population are
are the outcome of mutation breeding. collected and sown in upcoming season in mass selection. This
F I G U R E 1 (a) Field view and (b) seeds

of the browntop millet variety GPUBT-2.
F I G U R E 2 Small millet varieties released in

India employing various breeding methods.
6 NAGARAJA ET AL.
TABLE 3 Commercial small millets cultivars developed by Project Coordinating Unit, AICRP (Small millets), UAS Bangalore.
Year of Average
Cultivar Pedigree release yield (q/ha) Area of adaptation Special features
Finger millet
HR 374 EC 4840 IE 927 1975 20–25 Karnataka Profuse tillering, moderately
resistant to blast and viral disease,
compact tip incurved fingers and
pigmentation on all plant parts
HR 911 UAS1 IE 927 1986 25–28 Karnataka Erect plants, large and semi-
compact ears with tip incurved,
suitable for irrigated and rainfed
conditions
GPU 28 Indaf 5 (Indaf 9x IE 1998 35–40 Karnataka Medium tall plants, semi-compact
1012) ear with tip incurved fingers, broad
leaves with thick stem, highly
resistant to finger and neck blast
GPU 26 (I-5x I-9) IE 1012 2000 30–35 Karnataka Early duration, resistant to finger
and neck blast, suitable for early
kharif and late summer
GPU 45 GPU 26 L-5 2001 28–30 Gujarat, Karnataka, Madhya Early duration, suitable for late
Pradesh and Maharashtra kharif, semi-compact ear with tip
incurved fingers, highly resistant to
blast
GPU 48 GPU 26 L 5 2005 28–30 Karnataka Pigmentation on all plant parts and
(Ratna) highly resistant to blast
GPU 66 PR 202 GPU 28 2009 35–40 Karnataka Compact ears with tip incurved
fingers, light green narrow leaves
and bold grains, resistant to neck
and finger blast, suitable for late
kharif season
GPU 67 Selection from 2009 30–35 Chhattisgarh, Jharkhand, Semi-dwarf plants (85–90 cm) with
germplasm accession GE Karnataka, Maharashtra, Tamil decumbent plant type, medium
5331 Nadu and Uttarakhand compact ears with tip incurved
fingers
Foxtail millet
RA 118 Pureline selection 1976 15–16 Karnataka Long panicles and fodder yield
9.5 t/ha
K 221-1 Pureline selection 1976 25–30 Karnataka High yielding
SIA 3156 - 2012 20–25 Karnataka, Andhra Pradesh, Bihar, Highly responsive to nitrogenous
Gujarat, Madhya Pradesh, Tamil fertilizers
Nadu and Uttarakhand
GPUF 3 Selection from 2022 15–20 Karnataka Suitable for both kharif and
germplasm IC 479864 summer planting. Semi-compact
and oblong shaped inflorescence
with oval shaped yellow to
coloured seeds. Moderate resistant
to rust and leaf blight diseases
Proso millet
GPUP 21 GPUP 14 K1 2003 15–16 Karnataka and Tamil Nadu Non shattering, early maturity and
high fodder yield
GPUP 8 S7 L111 1999 12–15 Karnataka and Tamil Nadu Tall plants, broad flag leaf, large
compact panicles, bold grains and
resistant to brown spot disease
PMV 442 GPMS 109 GPMS 908 2019 18–20 Andhra Pradesh, Bihar, Karnataka, Short-duration, dwarf and non-
(GPUP 25) Tamil Nadu, Telangana and lodging variety. Grains are bold
Puducherry and grey in colour. Suitable for
rainfed conditions in kharif season
NAGARAJA ET AL. 7
TABLE 3 (Continued)
Year of Average
Cultivar Pedigree release yield (q/ha) Area of adaptation Special features
GPUP 28 GPUP 8 K 1 2021 17–20 Karnataka Suitable for kharif and summer
seasons. It has intermediate
compact with globose-
elliptic-shaped inflorescence. Grain
is oval shaped with golden yellow
colour. Moderately resistant to leaf
blight and resistant to brown spot
disease
Little millet
GPUL 6 JK 8 Peddasame 2021 15–20 Karnataka Suitable for kharif and summer
planting. Compact panicle with
arched shaped inflorescence. Grain
is oval shaped with dark brown
colour. Moderately resistant to leaf
blight and brown spot disease
Kodo millet
GPUK 3 Pureline selection from 1991 18–20 Tamil Nadu, Gujarat, Madhya Photo insensitive and high yielding
GPLM 826 Pradesh, Karnataka, Uttar Pradesh
and Maharashtra
Browntop millet
Hagari - 2021 18–20 Karnataka Suggested for Kharif sowing. Rich
Browntop 2 in Iron (70 mg/kg), zinc (42 mg/kg)
(HBr-2) and crude fibre (13%)
GPUBT 2 Selection from 2022 18–20 Karnataka Resistant to leaf blight with good
germplasm IC 617960 fodder yield
technique is also referred as phenotypic selection. An alternative pro- superior performance in comparison with the existing commercial cul-
cedure in mass selection includes harvest and grow the seeds from tivars are identified and released as a novel pureline variety. The rate
the best-looking plants separately and compare them. Often, mass at which the mutations are concentrated and gradual release of con-
selection is used to purify cultivars (landraces) and multiply those vari- cealed genetic variability through hybridization, natural cross pollina-
eties that are bred by pureline selection or pedigree breeding. How- tion, recombination and segregation determine favourable outcome of
ever, only marginal gains in plant performance in varieties like Gidda pure line selection. In finger millet and other small millets, pureline
Ragi and Hullubele of Karnataka, Saluchodi of Andhra Pradesh, Udu- selection has been commonly attempted. In India, it was Leslie
malpet Ragi and Guddapah Rajampet Ragi of Tamil Nadu and Murky C. Coleman, the second director of agriculture of Mysore State in Kar-
and Nangkatna of Sikkim are noticed that were developed by mass nataka, who initiated research on finger millet improvement in
selection. Often times, mass selection is of help for inter-varietal 1913 at Zonal Agricultural Research Station, V.C. Farm, Mandya, using
improvement for traits like grain yield, early maturity and disease pureline selection from indigenous cultivars like Hullubele and Gidda.
resistance (Harinarayana, 1986). In finger millet, ‘N40101’, an intro- In 1918, the first finger millet cultivar H-22 was developed by him,
duction from former Soviet Union was used to identify and select the subsequently identified numerous other cultivars such as K-1, R0870,
cultivar ‘KAT/PRO-1’ that produced 50% higher yield than the local ES-11, ES-13 CO-1 and H-1. In 1921 at Hathkamba (in Konkan),
cultivars, was developed by Kenya Agriculture Reforms and Innova- D-11, D-31 and A-16 purelines that were early, medium and late in
tion (KARI) to be grown in Kenya for commercial cultivation. It was maturity, respectively, were released. The other finger millet varieties
bred following mass selection with recurrent selection cycles for high developed and released in India are CO-2, CO-3, CO-7, CO-8, PLR-1,
yield. In proso millet, cultivar development and genetic improvement K-22, ES-11, RO-786, AKP6, VZM2 and Aruna. The pureline WWN-
have been obtained through the direct selection of promising acces- 25 is developed at Hill Millet Research Station, Navsari Agricultural
sions, similar to that in other small millets (Naylor et al., 2004). University, Waghai (Dangs) and released for cultivation in south and
middle Gujarat as Gujarat Navsari Nagli-7 (GNN-7) that is a white fin-
ger millet rich in calcium (468 mg/100 g) than the national check vari-
2.2.3 | Pureline selection ety VR-708 (398.0 mg/100 g) (Patil et al., 2016). In 1960, Rajendra
Prasad Agricultural University in Bihar developed Ram Cheena and
Progenies of the best-looking plants from a genetically variable popu- Sham Cheena using pureline selection. Subsequently, TNAU devel-
lation are evaluated morphologically for several years. Progenies with oped proso millet variety K2 by screening and selecting from local
8 NAGARAJA ET AL.
lines. In 2009, TNAU-145 and TNAU-151 were released from Coim- 4310 was obtained by crossing EC 1540 (low-yielding, white-grained
batore following pureline selection and that were resistant to rust and strain, high vitamin-13 content) with male parent EC 985 (high yield-
shoot fly damage (Gomashe, 2017). ing and brown grained) in Coimbatore, India (Wariar &
Divakaran, 1956).
In foxtail millet, hybridization-based pedigree selection became
2.2.4 | Hybridization popular in the 1970s, which is also the major breeding method
employed in many Chinese research institutes. Even so, in 1959, ‘Xin-
A popular breeding method employed in small millets is hybridization nongdong No. 2’ was the first hybridization-based foxtail millet vari-
to generate variability followed by selection in subsequent segregat- ety. Subsequently, ‘Sunrise’, ‘Huntsman’ and ‘Earlybird’ were the
ing population. The standardization of contact method of crossing in white-seeded proso millet varieties recognized from filial generations
finger millet prompted hybridization attempts in 1951. Hybridization of regional yield trials (Baltensperger, 1996, 2002). Most often, the
led to the development of Aruna, Poorna, Udaya, Annapurna and Cau- variety ‘Dawn’ was used as a parent to breed most of the proso millet
very in finger millet. In 1964, Dr. C. H. Lakshmanaiah attempted cultivars in Nebraska region (Lyon et al., 2008). In the United States, a
crosses between Indian varieties with African ecotypes to generate waxy proso millet cultivar ‘Plateau’ was developed by crossing waxy
novel variability. Potential African donor parents, namely, IE- 927, IE- donor parent PI 436626 with female parent ‘Huntsman’ (Santra
929, IE-980R, IR-810 and IE-902, were identified that were further et al., 2015). The Crop Breeding Institute of Heilongjiang Academy of
crossed with local cultivars such as Hallubele, K1, Annapurna, Purna, Agricultural Sciences, China, standardized proso millet hybridization
Cauvery, Shakti, Co-1 and Hamsa. His efforts led to the development protocol that is being followed and aided in the identification of
of 16 Indo-African varieties designated as ‘INDAF’ series 10 varieties including Longshu 16, the first Chinese proso millet vari-
(Lakshmanaiah, 1967). Among them, Indaf 9 was early, Co-9 and Indaf ety bred by the intraspecific hybridization technique
5 were mid-late and Indaf 1, Indaf 3, K5, Indaf 7, Indaf 8, Indaf 6, HR (Gomashe, 2017). Likewise, Nandini and Bhat (2019) identified a mod-
911 and Indaf 11 were late-maturing varieties. The productivity of ified hybridization method, that is, SMUASB in proso millet (Figure 3)
Indian exotic crosses was over 2500 kg/ha. This event gave a boost and little millet in India. In barnyard millet, inter-specific hybridization
to finger millet productivity with 50% increase in Karnataka between PRJ 1 and ER 72 cultivars of Echinochloa esculenta and Echi-
(Seetharam, 1989) and by 60% increase in Tamil Nadu (Nagarajan & nochloa frumentacea, respectively, produced vigorous, high tillering
Raveendran, 1983). Likewise, the first, white finger millet hybrid EC and culm branching hybrid free from smut disease, but seed setting
FIGURE 3 Modified hybridization method (SMUASB) in proso millet (Nandini & Bhat, 2019).
NAGARAJA ET AL. 9
failed due to sterility. The genetic variability spanning both the species intensive process that involves skilled manpower to maintain INFM
can be harnessed via hybridization ventures involving distant species 95001 has hampered its usage in hybridization programmes. To
(Sood et al., 2014). Millet breeders are encouraged to seek out cost- address this drawback, the finger millet variety GPU 28 was EMS trea-
effective, labour- and resource-efficient successful hybridization pro- ted to obtain novel partial sterile lines (PS1), in which partial sterility is
tocols that produce fully fertile hybrid seeds in other millets such as governed by single-recessive gene (Gowda et al., 2014); further, diver-
little millet, proso millet, kodo millet, barnyard millet, and browntop sification of PS lines with other genetic backgrounds (Figure 4a) along
millet, while standardising the hybridization protocols that are already with virescence markers was attempted (Figure 4b) (Manjappa
in use in finger millet and foxtail millet. To the best of the author's et al., 2023). The per cent seed set in crosses derived from PS1 ranged
knowledge. from 27.4% to 49.4% with an average of 40.1% and among 46 hybrids
the hybrid recovery ranged from 28.6 to 81.2% with an average of
56.7% (Manjappa, 2017). There is an ample scope to explore the use
2.2.5 | Male sterility of mutagens to induce male sterility in small millets especially consid-
ering that hybridization is difficult. Its practical efficacy is slightly due
Owing to the tiny floret size and incomprehensive floral biology of to differing levels of sterility/fertility across spatial locations and
small millets, simple and effortless successful hybridization is challeng- genetic backgrounds.
ing, thereby limiting the exploitation of hybrid vigour (Nagaraja Suanxi 28, a partial CMS foxtail millet line evolved from spontane-
et al., 2023; Nandini & Bhat, 2019). Therefore, identification of male- ous mutations in the landrace Suanpibai, is identified (Cui et al., 1979).
sterile lines which produces fertile hybrids upon crossing when used Upon being crossed with restorer lines, Suanxi 28 was able to produce
as parents in hybridization programme will be a feasible approach. hybrid seeds. Subsequently, herbicide-resistant cultivars were used as
Usage of male-sterile lines in hybrid seed production programme aids restorer lines for the development of herbicide-resistant foxtail millet
identification of novel recombinants with desirable combination of cultivars (Wang et al., 1996). Distant hybridization of Setaria verticil-
traits. Routinely, male-sterile lines have been identified and are being lata with foxtail millet generated a CMS line that is currently not being
utilized in seed production programmes in many commercial crops. used in hybrid seed production programmes for unknown reasons
Hot water treatment (52 C) of the finger millet inflorescence for (Zhu et al., 1991). Although several foxtail millet male-sterile lines
5 min (Raj et al., 1964) aided in converting into male-sterile line, have been developed, only partial genetic male-sterile (GMS)
besides the usage of fast neutrons (Mukuru et al., 1974) and ethyl lines have been used successfully in hybrid seed production (Diao &
methane sulphonate (EMS) mutagen (Gowda et al., 2014; Jia, 2017). Extensive screening of germplasm accessions or conversion
Gupta, 1999). The finger millet germplasm IE 3318 treated with EMS of a cultivar into male-sterile line is the need of the hour in little millet,
gave rise to genetic male-sterile line INFM 95001, in which the male proso millet, kodo millet, barnyard millet and browntop millet so that
sterility was governed by single-recessive gene. Upon being crossed they can be used as a parent in hybrid seed production programmes
with any homozygous (Ms1Ms1) parental line, INFM 95001 was able to develop non-lodging, abiotic and biotic stress-resistant cultivars
to produce fully fertile F1 progeny (Gupta et al., 1997). Labour- with high yielding and less seed shattering.
F I G U R E 4 (a) Finger millet panicle of

diversified partial sterile line and (b) appearance of
morphological virescence marker in partial
sterile line.
10 NAGARAJA ET AL.
2.2.6 | Mutation breeding greenhouse conditions (Konstantinov et al., 1991), while in barnyard
millet, a mutant lacking waxy protein was obtained from gamma irradi-
Plant mutation breeding, also referred as variation breeding, utilizes ation of low-amylase landrace ‘Nogehie’ that was stably inherited
physical radiations or chemical agents to incite spontaneous genetic (Hoshino et al., 2010). Further, a novel cultivar ‘Chojuromochi’ was
variation to develop novel and desirable cultivars. Mutation is the developed to popularize barnyard millet-based food products in Japan
source of numerous genetic variations and the driving force of evolu- (Goron & Raizada, 2015). Overall, mutation breeding has resulted in
tion. Mutation breeding in small millets began with irradiating finger the release of 13 small millets cultivars (finger millet 8, kodo millet
millet seeds using X-rays, resulting in development of Hagari-1 vari- 3 and little millet 2) in India.
ety, that was commercialized for cultivation in 1941 (Krishnaswami &
Ayyangar, 1941). Later on, the effects of gamma radiations and EMS
were deliberated in the finger millet cultivars Purna and Hamsa, ensu- 3 | S M A L L M I LL E T S B R E E D I N G T A R G E T S
ing this several worthy mutations were produced in Purna (Goud
et al., 1969, 1971). Consecutively, EMS and nitroso guanidine (NG)- Most commonly, almost all the varieties of small millets have been
treated finger millet seeds of AKP-7 resulted in identification of three bred by conventional breeding approach especially with a focus on
early maturing mutants (Sinha & Sahoo, 1971). A blast-resistant and improved yield levels. Despite significant accomplishments resulting
high-yielding finger millet mutant M21 developed by gamma irradiat- from conventional breeding methods, the drawbacks associated with
ing the cultivar HES 927 was identified by Shivashankar et al. (1973). them like genetic drag and erosion, reproductive hindrances and pro-
Hamsa, an early mutant of finger millet with increased finger number longed period it takes necessitate the need to identify resource-
and grain-bearing area, was obtained using gamma irradiations (Nayar efficient, quick and swift novel breeding methods that derive essential
et al., 1979), while Raveendran et al. (1982) obtained short-duration transformation needed in small millets' research and breeding. Addi-
and bold grain mutants from cultivars Sarada and MS2698, respec- tionally, with the current increasing popularity of small millets as a
tively, after treating with NG. Likewise, Gowda and Seetharam (1983) healthy diet option especially in the age of culinary enlightenment,
used gamma radiations on PR 202, HR-911, Indaf-8 and TNAU-294 this also aligns with the United Nations (UN) declaration of 2023 as
to identify several mutants that were dwarf, early maturing with high the International Year of Millets, to uplift the farming community by
capacity to tiller. The dwarf mutants of Annapurna and Cauvery varie- way of diversifying and embracing the cultivation of finger millet, fox-
ties of finger millet had relatively better harvest index and produced tail millet, little millet, proso millet, kodo millet, barnyard millet and
superior yield (Shivshankar & Ranganatha, 1983). Subsequently, browntop millet. This calls attention to expand the small millets breed-
attempts to identify desirable mutants from PR 202, Co 10, Indaf-9, ing objectives beyond increased yield levels, although yield is a com-
HR 24, IE 744 and GN1 upon being treated with gamma rays, EMS, plex and quantitatively inherited trait whose expression is dependent
methyl methane sulphonate (MMS) and diethyl sulphate (DES) were on numerous other traits. Consequently, genotype environment
initiated (Tikka, 1985). A white finger millet variety, Co 9 was treated interaction highly influences yield per se; therefore, estimating the
with low dosage of EMS to obtain high-yielding mutants stability of yield across multiple environments to ensure stable perfor-
(Devkota, 1987). The mutagenic efficiency in TRY 1 variety was inves- mance of the genotype with minimal yield loss before the cultivar is
tigated by Eswari et al. (2014), leading to the identification of desir- released for commercial cultivation is needed. The soft stalk of small
able mutants for plant height, tillers/plant, productive tillers, finger millets accompanied with increased number of tillers that result in
number, finger length and 1000 grain weight. In Zambia, the mutants higher biomass at maturity and the impact of various environmental
FMM165 and FMM175 developed using X-ray irradiation are regis- factors (Santra et al., 2019; Upadhyaya et al., 2015; Vetriventhan
tered in the FAO/IAEA Mutant Variety Database (https://mvd.iaea. et al., 2016) causes lodging. Yield loss that occur due to lodging is not
org). Therefore, mutation breeding was effectively used in finger millet yet been estimated till date. The weak culm diameter causes shatter-
for the development of early maturing types, generation of polygenic ing in small millets at harvest, resulting in poor yield levels
variability and development of complete/partial male-sterile lines. (Gomashe, 2017; Santra et al., 2019). Therefore, breeding cultivars
Physical, chemical and combinations of mutagens were used for this with strong stalk and culm diameter can result in realizing higher yield
purpose. It was in the 1970's that induced mutation breeding was ini- levels. Rajput and Santra (2016) detected quantitative trait loci (QTL)
tiated in foxtail millet. Suanxi 28, a partial CMS line (3% to 5% seed and flanking markers for seed shattering and lodging in proso millet
setting rate by self-fertilization) developed using mutation breeding that can further be utilized in marker-assisted breeding (MAB) to
was able to successfully set hybrid seeds when crossed with the develop shattering- and lodging-resistant cultivars.
restorer line (Cui et al., 1979). Use of radiation induced mutagenesis Owing to their adaptability to diverse agroecological conditions,
in breeding programmes has also produced some excellent foxtail mil- generally, small millets are less prone to the attack of insect pests and
let cultivars such as Chigu 4 (Li et al., 1990), Jingu 21 (Chen & pathogens. Often, poor cultivation practices and conducive environ-
Wei, 1992) and Longfu 93-076 (Yi et al., 2002). Jingu 21 was obtained mental conditions favour the development of disease or attack by
from γ-irradiation of Jinfen 52. However, considerable research on pests in endemic regions. Consequently, negligible crop protection
mutagens has not been undertaken in small millets. Smut-resistant measures are needed in small millets (Ravikesavan et al., 2022). How-
proso millet mutants expressed only 1.8% to 8.3% infection under ever, incidence of blast in finger millet (Nagaraja et al., 2007;
NAGARAJA ET AL. 11
Rao, 1990); blast and rust in foxtail millet (Vetriventhan et al., 2016); other hand, involves the use of such molecular marker data to
sheath blight, bacterial spot and head smut in proso millet; grain and enhance the effectiveness of various breeding activities, including
head smut and leaf spot diseases in barnyard millet; kodo millet; planning and execution of breeding programmes and improving selec-
and little millet (Upadhyaya et al., 2015) are reported. Likewise, the tion efficiency. It includes breeding methods, namely, marker-assisted
incidence of stem borer in finger millet and barnyard millet selection (MAS) and marker-assisted backcrossing (MABC), to transfer
(Upadhyaya et al., 2015) and shoot fly damage in proso millet, barn- oligogenic and polygenic controlled traits, MABC for gene pyramiding
yard millet, little millet and kodo millet are witnessed. This accentu- and marker-assisted recurrent selection (MARS) that can be
ates the need to identify pest- and disease-resistant sources that can attempted in small millets.
further be directly utilized as cultivar after several cycles of selection Omics-based approaches that are recurrently being employed in
or can be used as a parent in hybridization programme to develop most of the commercial crops can also be attempted in small millets.
pest- and disease-tolerant/resistant cultivars. Also, the tiny grain size Characterizing the candidate genes regulating various biological pro-
hampers milling recovery; thus, breeding for bold seeds would help cesses using functional genomics approaches like transcriptome-based
minimize post-harvest losses. Along with other breeding objectives, gene expression profiling shall contribute to the comprehensive infor-
next-generation sequencing technologies offer low-cost and high- mation about expression pattern of genes and functional polymor-
throughput analysis of plant genomes, allowing for better knowledge phism. As transcript abundance and expression does not directly
of genetic makeup and behaviour. Regular small millets breeding aims correlate with the protein/metabolome levels, and in order to study
include improving seed colour, nutrition profile, feed and fodder suit- the post-translational modifications that proteins/metabolomes
ability, flour quality, reducing anti-nutritional elements, and develop- undergo to perform any given function, proteomics and metabolomics
ing machine harvestable cultivars. Directing the focus on these studies are worthy (Sood et al., 2019).
breeding objectives shall result in improved rate of genetic advance- A reference genome of any crop of small millets shall serve as an
ment in small millets by way of harnessing their genetic potential. asset to understand important and adaptive phenotypes, yet it is
The approach to address the above-mentioned breeding objectives becoming clear that significant advances in breeding-improved varie-
in millets is constrained not just by difficulties in hybridization tech- ties are only possible when the genomic variants are known, thus
niques but also due to scanty genomic resources. The recent genomic requiring a population of genomes and potentially a pangenome
advances, availability of wide choice of genomic tools and technologies (Chapman et al., 2022). The discovery and mapping of genome-wide
and the reduced sequencing cost have boosted genome-scale research allelic variation is aided by genome sequence information. Besides,
to hasten crop improvement especially in the popularly cultivated com- adopting efficient family-based linkage designs/large diversity panels,
mercial crops that can also be efficiently utilized in breeding millets. multi-omics assays and high-throughput phenotyping (HTP) platforms
They have been of help to improve the understanding of genome shall assist in developing genome–phenome maps.
structure and function and decipher the genetic behaviour of important GAB of millets has been limited due to the lack of molecular
traits. Extensive genomic studies and identification of novel genetic markers linked to traits of interest, but the recent recognition of the
determinants controlling beneficial traits in millets can be systematically importance of millets and the availability of advanced genomic tools
mined to design breeding strategies to overcome the stagnation of and technologies provide a solution for this. The high-density single-
yield while maintaining the nutrient density and overcome the agro- nucleotide polymorphism (SNP) data obtained from population-wide
climatic and post-harvest challenges using the novel genome-based genome sequencing can be used to identify the genetic basis of adap-
breeding methods (Muthamilarasan et al., 2019). Numerous improved tive traits through genome-wide association studies (GWAS) and
crop products developed using genomic innovations are available for selection analyses. From this, genetic markers can be designed
cultivation in farmers' fields but none in small millets. and used for MAB. Extensive genomic studies and identification of
novel genetic determinants controlling beneficial traits in millets can
be systematically mined to design breeding strategies to overcome
3.1 | Genomics-based breeding in small millets the stagnation of yield while maintaining the nutrient density and
overcome the agro-climatic and post-harvest challenges. Extended
With the advent of next-generation sequencing technologies offer population genomics, for example, using reduced representation tech-
low-cost and high-throughput analysis of plant genomes, that enable nologies, can be used to link genetic markers to agronomic traits,
for better knowledge of genetic makeup and behaviour. Plant whole- without knowing the precise genetic basis of the trait. Markers linked
genome sequencing, transcriptome sequencing and exome sequencing to several diverse agronomic traits in foxtail millet have been identi-
will allow discovering agronomically important genes that regulate fied using these approaches, for example, coloration, leaf size and
yield and tolerance to biotic and abiotic stresses. The data generated shape, grain yield and weight and flowering time (Jaiswal et al., 2019;
from genome and transcriptome sequencing in a cost-effective way Upadhyaya et al., 2015). Although genomic resources have been iden-
enable profiling of nucleotide variation and discover regulatory tified in small millets for various traits (Table 4), their utility in marker
sequences and their relative positions and large-scale discovery of assisted breeding is limited. However, they can be used to decipher
molecular markers that can be further used in genomics-assisted trait behaviour, introduce desired QTLs in a superior yielding cultivar
breeding (GAB) (Varshney et al., 2021). Molecular breeding, on the and to breed novel small millet cultivars.
12 NAGARAJA ET AL.
TABLE 4 Genomic resources available in small millets.
Breeding material No. of genes annotated/

Trait used Molecular marker system QTLs and SNPs identified Reference
Finger millet
Calcium content - Anchored expressed sequence 14 markers Kumar et al. (2015)
tagged–simple sequence repeats
(EST SSRs)
Calcium content - Anchored SSRs 5 markers Nirgude et al. (2014)
Drought responsive - Whole-genome sequencing 2866 SNPs Hittalmani et al. (2017)
R-genes - Whole-genome sequencing 1766 SNPs Hittalmani et al. (2017)
Calcium transport and - Whole-genome sequencing 330 SNPs Hittalmani et al. (2017)
accumulation
- - Whole-genome sequencing 62,348 SNPs Hatakeyama et al. (2018)
Calcium 113 germplasm Association mapping 9 QTLs Kumar et al. (2015)
Basal tiller number, days to 190 germplasm SSR 46 markers Babu, Agrawal, Pandey,
50% flowering, flag leaf Jaiswal, and Kumar (2014)
blade width and plant height
Finger blast 190 germplasm SSR 104 markers and 4 QTLs Babu, Dinesh, Agrawal,
Sood, et al. (2014)
Tryptophan 190 germplasm SSR 120 markers and 2 QTL Babu, Agrawal, Pandey,
Protein 1 QTL and Kumar (2014)
Productive tillers, seed yield, 128 germplasm SSR 87 markers and 7 QTLs Ramakrishnan et al.
leaf blast resistance and (2016)
number of tillers
Root dry weight, shoot dry - SSR 4 QTLs Ramakrishnan et al.
weight and root length (2017)
Phosphate transporter 4 varieties Expression studies 4 genes Pudake et al. (2017)
Salinity stress - Expression studies 1 gene Ramegowda et al. (2012)
Abiotic stress response GPU-28 Expression studies 2 genes Babitha et al. (2015)
Drought tolerance - Expression studies 1 gene Nagarjuna et al. (2016)
Drought stress GPU-28 Expression studies 1 gene Parvathi and Nataraja
(2017)
Saline and heat stresses - Expression studies 1 gene Ramakrishna et al. (2018)
Foxtail millet
Biofuel - Whole-genome sequencing 38,801 SNPs Zhang et al. (2012)
Putative proteins - EST SSRs 11 Jia, Shi, et al. (2007)
Basal branching F3 families Restriction fragment length 4 QTLs Doust et al. (2004)
polymorphism (RFLP)
Dehydration tolerance 107 cultivars Biochemical marker 1 gene Lata et al. (2011)
Salt tolerance 2 varieties Expression studies 1 gene Sreenivasulu et al. (2004)
Herbicide tolerances 11 genotypes Expression studies 1 gene Jia, Yuan, et al. (2007)
Drought tolerance 5 genotypes Expression studies 1 gene Zhang et al. (2007)
Aluminium tolerance 1 genotype Expression studies 1 gene Zhao et al. (2009)
Drought and heat tolerance 9 genotypes Genomic in situ hybridization 1 gene Wang et al. (2009)
(GISH)
Salinity and drought 1 genotype Expression studies 1 gene Mishra et al. (2012)
tolerance
Nutrient Association GWAS 10 marker trait association Jaiswal et al. (2019)
mapping panel
Transferability and 96 genotypes Intron length polymorphic 5123 markers Muthamilarasan et al.
phylogenetics marker (2014)
NAGARAJA ET AL. 13
TABLE 4 (Continued)
Breeding material No. of genes annotated/

Trait used Molecular marker system QTLs and SNPs identified Reference
Little millet
- 25 genotypes SSR 4443 EST SSRs Desai et al. (2021)
- 36 Random amplified polymorphic 60 Tiwari et al. (2018)
DNA (RAPD)
Proso millet
Protein-coding genes and 132 recombinant Whole-genome sequencing 222,081 SNPs Zou et al. (2019)
microRNA genes inbred lines (RILs)
Diversity studies 50 accessions SSR 25 markers Cho et al. (2010)
Diversity studies 96 SSR 16 markers Hunt et al. (2011)
Diversity studies 8 accessions SSR 100 Rajput et al. (2017)
Diversity studies 88 SSR 67 markers Liu et al. (2016)
Morpho-agronomic traits 833 GBS 18 QTLs Rajput and Santra (2016)
Stress tolerance 2 genotypes De novo assembly 32,216 SSRs and 400,000 Yue et al. (2016)
SNP loci
Barnyard millet
Waxy trait - SNP 3 genes Ishikawa et al. (2013)
Anthocyanin pigment - SSR 1 gene Renganathan et al. (2019)
Discriminating SNPs 89 genotypes GBS 2579 SNPs Wallace et al. (2015)
Precise and rapid alterations in plant genomes are possible via plants have limited its usage in small millets breeding. A combination
advances in genome editing techniques. In recent years, genetic engi- of these approaches shall provide the necessary support to develop
neering has been widely used to elucidate gene function and for crop improved small millets varieties.
improvement. Umpteen genetic transformation studies have been
attempted in major cereals like rice, wheat and maize, but lagging in
small millets (Ceasar & Ignacimuthu, 2009; Vetriventhan et al., 2020). 4 | FU T U R E C H A L L E NG E S A N D
However, quite a few attempts to standardize transformation proto- PROSPECTS
cols and development of transgenic plants expressing desirable genes
have been initiated (Vetriventhan et al., 2020), Liu et al. (2005) With potential role of small millets in food and nutritional security, it
reported biolistic method and Agrobacterium-mediated transformation is essential to tap their nutritional profile for our day-to-day dietary
in finger millet and foxtail millet, respectively, for the first time. Subse- requirements to lead a healthy lifestyle. To achieve this goal, primarily,
quently, emphasis was given to develop transgenics for leaf blast the challenges to breed small millets need to be addressed. Besides
resistance (Ignacimuthu & Ceasar, 2012; Latha et al., 2005), drought understanding the flower structure and size of small millets, determin-
and salinity tolerance (Hema et al., 2014), herbicide resistance (Bayer ing their pollen viability, stigma receptivity and time of pollination is
et al., 2014) and biofortification of zinc (Ramegowda et al., 2013). In essential to standardize hybridization techniques in which desirable
foxtail millet, drought and salinity tolerant (Li et al., 2014; Li parents obtained by screening the available germplasm accessions are
et al., 2017; Pan et al., 2016; Wang et al., 2014) and characterization crossed to derive superior trait-specific genetic recombinants not
of phosphate transporters studies are initiated (Ceasar et al., 2017). existing in the available gene pool, thereby exploiting hybrid vigour.
Very few genetic transformation studies are attempted in barnyard For the ease of seed multiplication, identification of male-sterile
millet. There are no reports on little millet, proso millet, kodo millet sources that can be used as parents in hybridization programmes shall
and browntop millet. But in vitro regeneration protocols in little millet, serve as a cost-effective strategy. Use of mutagens to achieve the
proso millet, kodo millet and barnyard millet have been initiated. In mentioned objectives shall boost small millets breeding activities. Fur-
2011, ‘Regent’, a proso millet cultivar, was developed using anther ther, with the availability of next-generation sequencing tools at
culture technique; subsequent selections were attempted for produc- affordable price, genome sequencing to identify molecular markers
tivity and grain-quality parameters. It is medium duration (95– that are of help in GAB, transcriptome profiling to decipher gene
105 days), resistant to lodging, shattering, majority of races of smut, structure and function, proteome profiling that aids in identification of
melanosis disease and high yielding. Due to the frequent occurrence peptides and proteins governing desirable traits and metabolome pro-
of albinism especially in monocots, difficulties involved in excision of filing to dissect the genotype–phenotype relationship in response to
anthers accompanied with reduced success rate of obtaining haploid the various biotic and abiotic stresses is encouraged. Additionally,
14 NAGARAJA ET AL.
breeding objectives centred to develop cultivars with dual purpose, Babu, B. K., Dinesh, P., Agrawal, P. K., Sood, S., Chandrashekara, C., &
high climate resilient, biofortified with increased level of bioavailability Bhatt, J. C. (2014). Comparative genomics and association mapping
approaches for blast resistant genes in finger millet using SSRs. PLoS
of nutrients and resistant to biotic and abiotic stresses shall be incen-
ONE, 9, e99182. https://doi.org/10.1371/journal.pone.0099182
tivized. To achieve food and nutritional security in the changing cli- Baltensperger, D. D. (1996). Foxtail and proso millet. In J. Janick (Ed.), Pro-
matic conditions, it is the need of the hour to promote the cultivation gress in new crops (pp. 182–190). ASHS Press.
of small millets and include them in our diets, besides ensuring our Baltensperger, D. D. (2002). Progress with proso, pearl and other millets.
In J. Janick & A. Whipkey (Eds.), Trends in new crops and new uses
focus on research and development in small millets.
(pp. 100–103). ASHS Press.
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Received: 27 January 2024 Revised: 6 June 2024 Accepted: 9 July 2024

Panorama of small millets breeding: A review

T. E. Nagaraja 1 | S. Gazala Parveen 1 | J. Meenakshi 1 | S. Murtujasab 1 |

1 | I N T RO DU CT I O N minor millets/small millets (Gupta, 2006).Millets thrive well in low

TABLE 1 Details of small millets discussed in this review.

Note: -: Data are not available.

TABLE 2 Nutrient composition of small millets (per 100 g).

Note: -: Data are not available.

F I G U R E 1 (a) Field view and (b) seeds

F I G U R E 2 Small millet varieties released in

F I G U R E 4 (a) Finger millet panicle of

TABLE 4 Genomic resources available in small millets.

Breeding material No. of genes annotated/

Breeding material No. of genes annotated/

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