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RESEARCH ARTICLE

Exploring morpho-physiological, biochemical, and molecular strategies of finger millet (Eleusine coracana) for drought stress tolerance: a review

Srikanth K https://orcid.org/0009-0002-8604-6415 A , Asish Binodh A * , Ravikesavan Rajasekaran https://orcid.org/0000-0002-4250-5854 A , Alagarsamy Senthil B and Narayanan Manikanda Boopathi C
+ Author Affiliations
- Author Affiliations

A Centre for Plant Breeding and Genetics, Tamil Nadu Agricultural University, Coimbatore 641003, India.

B Department of Crop Physiology, Tamil Nadu Agricultural University, Coimbatore 641003, India.

C Department of Plant Biotechnology, Tamil Nadu Agricultural University, Coimbatore 641003, India.

* Correspondence to: akbinodh@gmail.com

Handling Editor: Enrico Francia

Crop & Pasture Science 76, CP25028 https://doi.org/10.1071/CP25028
Submitted: 25 January 2025  Accepted: 2 April 2025  Published: 23 April 2025

© 2025 The Author(s) (or their employer(s)). Published by CSIRO Publishing

Abstract

Finger millet (Eleusine coracana (L.) Gaertn.) is a climate-resilient C4 cereal crop with exceptional adaptability to arid and semi-arid regions. Its unique morpho-physiological, biochemical, and molecular mechanisms contribute to its high tolerance to drought, making it a valuable model for sustainable agriculture and breeding programmes. This review explored the drought resilience strategies of finger millet, focusing on its genetic traits, physiological adaptations, and molecular mechanisms. Additionally, the study examines its potential for guiding resilient cropping systems and improving drought tolerance in major crops. A comprehensive review of recent research was conducted, analysing morpho-physiological traits such as root architecture, stomatal control, and biochemical responses, including osmolyte accumulation and antioxidant activities. Molecular studies identifying stress-responsive genes and transcriptomic pathways were also evaluated. Finger millet exhibits high water use efficiency, robust root systems, and adaptive morphological traits that enhance drought resilience. Biochemical responses, such as proline and soluble sugar accumulation, mitigate osmotic stress and oxidative damage. Molecular studies identified key drought-responsive genes (EcDehydrin7, EcNAC67, EcbZIP60) and revealed syntenic relationships with Poaceae species, facilitating gene transfer for breeding. Finger millet’s diverse genetic traits and stress-tolerance mechanisms make it an essential resource for improving drought tolerance in major crops and developing climate-smart agriculture. The insights from finger millet can guide breeding programmes and agricultural practices to enhance global food security in the face of climate change.

Keywords: antioxidant enzymes, biochemical mechanisms, drought, drought-responsive genes, finger millet, millets, morphological adaptations, physiological responses.

References

Abd Allah AA, Badawy SA, Zayed BA, El-Gohary AA (2010) The role of root system traits in the drought tolerance of rice (Oryza sativa L.). Journal of Plant Production 1(4), 621-631.
| Crossref | Google Scholar |

Ahluwalia O, Singh PC, Bhatia R (2021) A review on drought stress in plants: implications, mitigation and the role of plant growth promoting rhizobacteria. Resources, Environment and Sustainability 5, 100032.
| Crossref | Google Scholar |

Ahmad S, Ahmad R, Ashraf MY, Ashraf M, Waraich EA (2009) Sunflower (Helianthus annuus L.) response to drought stress at germination and seedling growth stages. Pakistan Journal of Botany 41(2), 647-654.
| Google Scholar |

Ahmad P, Jaleel CA, Salem MA, Nabi G, Sharma S (2010) Roles of enzymatic and nonenzymatic antioxidants in plants during abiotic stress. Critical Reviews in Biotechnology 30(3), 161-175.
| Crossref | Google Scholar | PubMed |

Akram NA, Shafiq F, Ashraf M (2017) Ascorbic acid – a potential oxidant scavenger and its role in plant development and abiotic stress tolerance. Frontiers in Plant Science 8, 613.
| Crossref | Google Scholar |

Antony Ceasar S, Maharajan T, Ajeesh Krishna TP, Ramakrishnan M, Victor Roch G, Satish L, Ignacimuthu S (2018) Finger millet [Eleusine coracana (L.) Gaertn.] improvement: current status and future interventions of whole genome sequence. Frontiers in Plant Science 9, 1054.
| Crossref | Google Scholar |

Ashoka P, Raut D, Sudeepthi B, Gawande KN, Reddy GSV, Padhan SR, Panigrahi CK (2023) Millet’s role as a climate resilient staple for future food security: a review. International Journal of Environment and Climate Change 13(11), 4542-4552.
| Crossref | Google Scholar |

Ashraf M, Foolad MR (2007) Roles of glycine betaine and proline in improving plant abiotic stress resistance. Environmental and Experimental Botany 59(2), 206-216.
| Crossref | Google Scholar |

Ashraf M, Harris PJC (2013) Photosynthesis under stressful environments: an overview. Photosynthetica 51, 163-190.
| Crossref | Google Scholar |

Babitha KC, Ramu SV, Nataraja KN, Sheshshayee MS, Udayakumar M (2015) EcbZIP60, a basic leucine zipper transcription factor from Eleusine coracana L. improves abiotic stress tolerance in tobacco by activating unfolded protein response pathway. Molecular Breeding 35, 181.
| Crossref | Google Scholar |

Backiyalakshmi C, Vetriventhan M, Deshpande S, Babu C, Allan V, Naresh D, Gupta R, Azevedo VCR (2021) Genome-wide assessment of population structure and genetic diversity of the global finger millet germplasm panel conserved at the ICRISAT Genebank. Frontiers in Plant Science 12, 692463.
| Crossref | Google Scholar |

Badr A, El-Shazly HH, Tarawneh RA, Börner A (2020) Screening for drought tolerance in maize (Zea mays L.) germplasm using germination and seedling traits under simulated drought conditions. Plants 9(5), 565.
| Crossref | Google Scholar |

Bai J, Gong C-M, Chen K, Kang H-M, Wang G (2009) Examination of antioxidative system’s responses in the different phases of drought stress and during recovery in desert plant Reaumuria soongorica (Pall.) Maxim. Journal of Plant Biology 52, 417-425.
| Crossref | Google Scholar |

Bandyopadhyay T, Muthamilarasan M, Prasad M (2017) Millets for next generation climate-smart agriculture. Frontiers in Plant Science 8, 1266.
| Crossref | Google Scholar |

Bartwal A, Arora S (2017) Drought stress-induced enzyme activity and mdar and apx gene expression in tolerant and susceptible genotypes of Eleusine coracana (L.). In Vitro Cellular & Developmental Biology-Plant 53, 41-49.
| Crossref | Google Scholar |

Bartwal A, Pande A, Sharma P, Arora S (2016) Intervarietal variations in various oxidative stress markers and antioxidant potential of finger millet (Eleusine coracana) subjected to drought stress. Journal of Environmental Biology 37(4), 517-522.
| Google Scholar | PubMed |

Behera MK (2017) Assessment of the state of millets farming in India. MOJ Ecology & Environmental Science 2(1), 16-20.
| Crossref | Google Scholar |

Behnamnia M, Kalantari KM, Rezanejad F (2009) Exogenous application of brassinosteroid alleviates drought-induced oxidative stress in Lycopersicon esculentum L. General and Applied Plant Physiology 35(1–2), 22-34.
| Google Scholar |

Bhatt D, Negi M, Sharma P, Saxena SC, Dobriyal AK, Arora S (2011) Responses to drought induced oxidative stress in five finger millet varieties differing in their geographical distribution. Physiology and Molecular Biology of Plants 17, 347-353.
| Crossref | Google Scholar | PubMed |

Bibi A, Sadaqat HA, Akram HM, Mohammed MI (2010) Physiological markers for screening sorghum (Sorghum bicolor) germplasm under water stress condition. International Journal of Agriculture & Biology 12(3), 451-455.
| Google Scholar |

Bohnert HJ, Jensen RG (1996) Strategies for engineering water-stress tolerance in plants. Trends in Biotechnology 14(3), 89-97.
| Crossref | Google Scholar |

Bolouri-Moghaddam MR, Le Roy K, Xiang L, Rolland F, Van den Ende W (2010) Sugar signalling and antioxidant network connections in plant cells. The FEBS Journal 277(9), 2022-2037.
| Crossref | Google Scholar | PubMed |

Bwalya EC, Marzougui S, Mwangi E, Wooseon C, Lee M-C (2020) Genetic diversity analysis and population structure of some African and Asian Finger Millet (Eleusine coracana L.) accessions using expressed sequence tags–simple sequence repeat (EST-SSR) markers. International Journal of Environment, Agriculture and Biotechnology 5(3), 643-646.
| Crossref | Google Scholar |

Cao H-X, Sun C-X, Shao H-B, Lei X-T (2011) Effects of low temperature and drought on the physiological and growth changes in oil palm seedlings. African Journal of Biotechnology 10(14), 2630-2637.
| Crossref | Google Scholar |

Cattivelli L, Rizza F, Badeck F-W, Mazzucotelli E, Mastrangelo AM, Francia E, Marè C, Tondelli A, Stanca AM (2008) Drought tolerance improvement in crop plants: an integrated view from breeding to genomics. Field Crops Research 105(1–2), 1-14.
| Crossref | Google Scholar |

Chaniago I, Syarif A, Riviona P (2017) Sorghum seedling drought response: in search of tolerant genotypes. International Journal on Advanced Science, Engineering and Information Technology 7(3), 892-897.
| Crossref | Google Scholar |

Chinnusamy V, Jagendorf A, Zhu J-K (2005) Understanding and improving salt tolerance in plants. Crop Science 45(2), 437-448.
| Crossref | Google Scholar |

Chivenge P, Mabhaudhi T, Modi AT, Mafongoya P (2015) The potential role of neglected and underutilised crop species as future crops under water scarce conditions in Sub-Saharan Africa. International Journal of Environmental Research and Public Health 12(6), 5685-5711.
| Crossref | Google Scholar | PubMed |

Comas LH, Becker SR, Cruz VMV, Byrne PF, Dierig DA (2013) Root traits contributing to plant productivity under drought. Frontiers in Plant Science 4, 442.
| Crossref | Google Scholar |

Das K, Roychoudhury A (2014) Reactive oxygen species (ROS) and response of antioxidants as ROS-scavengers during environmental stress in plants. Frontiers in Environmental Science 2, 53.
| Crossref | Google Scholar |

Debieu M, Sine B, Passot S, Grondin A, Akata E, Gangashetty P, Vadez V, Gantet P, Foncéka D, Cournac L, Hash CT, Kane NA, Vigouroux Y, Laplaze L (2018) Response to early drought stress and identification of QTLs controlling biomass production under drought in pearl millet. PLoS ONE 13(10), e0201635.
| Crossref | Google Scholar |

Delauney AJ, Verma DPS (1993) Proline biosynthesis and osmoregulation in plants. The Plant Journal 4(2), 215-223.
| Crossref | Google Scholar |

Demiral T, Türkan I (2004) Does exogenous glycinebetaine affect antioxidative system of rice seedlings under NaCl treatment? Journal of Plant Physiology 161(10), 1089-1100.
| Crossref | Google Scholar | PubMed |

de Wet JMJ (2006) Eleusine coracana (L.) Gaertn. Record from Protabase. PROTA (Plant Resources of Tropical Africa/Ressources Végétales de l’Afrique Tropicale) Wageningen, Netherlands.

Dichio B, Xiloyannis C, Angelopoulos K, Nuzzo V, Bufo SA, Celano G (2003) Drought-induced variations of water relations parameters in Olea europaea. Plant and Soil 257, 381-389.
| Crossref | Google Scholar |

Dien DC, Thu TTP, Moe K, Yamakawa T (2019) Proline and carbohydrate metabolism in rice varieties (Oryza sativa L.) under various drought and recovery conditions. Plant Physiology Reports 24, 376-387.
| Crossref | Google Scholar |

Doorenbos J, Kassam AH, Bentvelsen C, Uittenbogaard G (1979) Yield response to water. In ‘Irrigation and agricultural development. Vol. 33’. (Ed. SS Johl) pp. 257–280. (Pergamon) 10.1016/B978-0-08-025675-7.50021-2

Duque AS, de Almeida AM, da Silva AB, da Silva JM, Farinha AP, Santos D, Fevereiro P, de Sousa Araújo S (2013) Abiotic stress responses in plants: unraveling the complexity of genes and networks to survive. In ‘Abiotic stress-plant responses and applications in agriculture’. (Eds K Vahdati, C Leslie) pp. 49–101. (InTech)

Evers D, Lefèvre I, Legay S, Lamoureux D, Hausman J-F, Rosales ROG, Marca LRT, Hoffmann L, Bonierbale M, Schafleitner R (2010) Identification of drought-responsive compounds in potato through a combined transcriptomic and targeted metabolite approach. Journal of Experimental Botany 61(9), 2327-2343.
| Crossref | Google Scholar | PubMed |

Faisal S, Mujtaba SM, Khan MA, Mahboob W (2017) Morpho-physiological assessment of wheat (Triticum aestivum L.) genotypes for drought stress tolerance at seedling stage. Pakistan Journal of Botany 49(2), 445-452.
| Google Scholar |

Fracasso A, Magnanini E, Marocco A, Amaducci S (2017) Real-time determination of photosynthesis, transpiration, water-use efficiency and gene expression of two Sorghum bicolor (Moench) genotypes subjected to dry-down. Frontiers in Plant Science 8, 932.
| Crossref | Google Scholar |

Fraser TE, Silk WK, Rost TL (1990) Effects of low water potential on cortical cell length in growing regions of maize roots. Plant Physiology 93(2), 648-651.
| Crossref | Google Scholar | PubMed |

Fridovich I (1981) Superoxide radical and superoxide dismutases. In ‘Oxygen and living processes: an interdisciplinary approach’. (Ed. DL Gilbert) pp. 250–272. (Springer) 10.1007/978-1-4612-5890-2_13

Gajewska E, Skłodowska M (2008) Differential biochemical responses of wheat shoots and roots to nickel stress: antioxidative reactions and proline accumulation. Plant Growth Regulation 54, 179-188.
| Crossref | Google Scholar |

Galeano E, Vasconcelos TS, Novais de Oliveira P, Carrer H (2019) Physiological and molecular responses to drought stress in teak (Tectona grandis L.f.). PLoS ONE 14(9), e0221571.
| Crossref | Google Scholar |

Ge T-D, Sui F-G, Bai L-P, Lu Y-Y, Zhou G-S (2006) Effects of water stress on the protective enzyme activities and lipid peroxidation in roots and leaves of summer maize. Agricultural Sciences in China 5(4), 291-298.
| Crossref | Google Scholar |

Gill SS, Tuteja N (2010) Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants. Plant Physiology and Biochemistry 48(12), 909-930.
| Crossref | Google Scholar | PubMed |

Gimode D, Odeny DA, de Villiers EP, Wanyonyi S, Dida MM, Mneney EE, Muchugi A, Machuka J, de Villiers SM (2016) Identification of SNP and SSR markers in finger millet using next generation sequencing technologies. PLoS ONE 11(7), e0159437.
| Crossref | Google Scholar |

Gomes FP, Oliva MA, Mielke MS, Almeida A-AF, Aquino LA (2010) Osmotic adjustment, proline accumulation and cell membrane stability in leaves of Cocos nucifera submitted to drought stress. Scientia Horticulturae 126(3), 379-384.
| Crossref | Google Scholar |

Gomez-Zavaglia A, Mejuto JC, Simal-Gandara J (2020) Mitigation of emerging implications of climate change on food production systems. Food Research International 134, 109256.
| Crossref | Google Scholar |

Gupta SM, Arora S, Mirza N, Pande A, Lata C, Puranik S, Kumar J, Kumar A (2017) Finger millet: a “certain” crop for an “uncertain” future and a solution to food insecurity and hidden hunger under stressful environments. Frontiers in Plant Science 8, 643.
| Crossref | Google Scholar |

Hadebe ST, Modi AT, Mabhaudhi T (2017) Drought tolerance and water use of cereal crops: a focus on sorghum as a food security crop in sub-Saharan Africa. Journal of Agronomy and Crop Science 203(3), 177-191.
| Crossref | Google Scholar |

Hanson AD, Nelsen CE, Everson EH (1977) Evaluation of free proline accumulation as an index of drought resistance using two contrasting barley cultivars. Crop Science 17(5), 720-726.
| Crossref | Google Scholar |

Hare PD, Cress WA (1997) Metabolic implications of stress-induced proline accumulation in plants. Plant Growth Regulation 21, 79-102.
| Crossref | Google Scholar |

Hellal FA, El-Shabrawi HM, Abd El-Hady M, Khatab IA, El-Sayed SAA, Abdelly C (2018) Influence of PEG induced drought stress on molecular and biochemical constituents and seedling growth of Egyptian barley cultivars. Journal of Genetic Engineering and Biotechnology 16(1), 203-212.
| Crossref | Google Scholar | PubMed |

Hittalmani S, Mahesh HB, Shirke MD, Biradar H, Uday G, Aruna YR, Lohithaswa HC, Mohanrao A (2017) Genome and Transcriptome sequence of Finger millet (Eleusine coracana (L.) Gaertn.) provides insights into drought tolerance and nutraceutical properties. BMC Genomics 18(1), 465.
| Crossref | Google Scholar |

Hosseini SA, Réthoré E, Pluchon S, Ali N, Billiot B, Yvin J-C (2019) Calcium application enhances drought stress tolerance in sugar beet and promotes plant biomass and beetroot sucrose concentration. International Journal of Molecular Sciences 20(15), 3777.
| Crossref | Google Scholar |

Host Antony David R, Ramakrishnan M, Maharajan T, BarathiKannan K, Atul Babu G, Daniel MA, Agastian P, Antony Caesar S, Ignacimuthu S (2021) Mining QTL and genes for root traits and biochemical parameters under vegetative drought in South Indian genotypes of finger millet (Eleusine coracana (L.) Gaertn) by association mapping and in silico comparative genomics. Biocatalysis and Agricultural Biotechnology 32, 101935.
| Crossref | Google Scholar |

Hu W, Huang Y, Loka DA, Bai H, Liu Y, Wang S, Zhou Z (2020) Drought-induced disturbance of carbohydrate metabolism in anthers and male abortion of two Gossypium hirsutum cultivars differing in drought tolerance. Plant Cell Reports 39, 195-206.
| Crossref | Google Scholar | PubMed |

Huang J, Ji M, Xie Y, Wang S, He Y, Ran J (2016) Global semi-arid climate change over last 60 years. Climate Dynamics 46, 1131-1150.
| Crossref | Google Scholar |

Hussain MI, Muscolo A, Farooq M, Ahmad W (2019) Sustainable use and management of non-conventional water resources for rehabilitation of marginal lands in arid and semiarid environments. Agricultural Water Management 221, 462-476.
| Crossref | Google Scholar |

Jung J-S, Muhammad Z, Lee K-W, Mun J-Y, Park H-S, Kim Y-J, Kim W-H, Lee S-H, Lee S-H (2015) Effects of polyethylene glycol-induced water stress on the physiological and biochemical responses of different Sorghum genotypes. IGC Proceedings (1989-2023). 4. https://uknowledge.uky.edu/igc/23/4-1-2/4

Kapoor D, Bhardwaj S, Landi M, Sharma A, Ramakrishnan M, Sharma A (2020) The impact of drought in plant metabolism: how to exploit tolerance mechanisms to increase crop production. Applied Sciences 10(16), 5692.
| Crossref | Google Scholar |

Kato Y, Kamoshita A, Yamagishi J (2008) Preflowering abortion reduces spikelet number in upland rice (Oryza sativa L.) under water stress. Crop Science 48(6), 2389-2395.
| Crossref | Google Scholar |

Khaleghi A, Naderi R, Brunetti C, Maserti BE, Salami SA, Babalar M (2019) Morphological, physiochemical and antioxidant responses of Maclura pomifera to drought stress. Scientific Reports 9(1), 19250.
| Crossref | Google Scholar |

Khatoon H, Singh V (2016) Impact of water stress on physiological and biochemical parameters of finger millet (Eleusine coracana L.). Research in Environment and Life Sciences 9(12), 1474-1477.
| Google Scholar |

Kholová J, Hash CT, Kočová M, Vadez V (2011) Does a terminal drought tolerance QTL contribute to differences in ROS scavenging enzymes and photosynthetic pigments in pearl millet exposed to drought? Environmental and Experimental Botany 71(1), 99-106.
| Crossref | Google Scholar |

Kishor PBK, Sangam S, Amrutha RN, Laxmi PS, Naidu KR, Rao KRSS, Rao S, Reddy KJ, Theriappan P, Sreenivasulu N (2005) Regulation of proline biosynthesis, degradation, uptake and transport in higher plants: its implications in plant growth and abiotic stress tolerance. Current Science 88, 424-438.
| Google Scholar |

Köşkeroğlu S, Tuna AL (2010) The investigation on accumulation levels of proline and stress parameters of the maize (Zea mays L.) plants under salt and water stress. Acta Physiologiae Plantarum 32, 541-549.
| Crossref | Google Scholar |

Kotapati KV, Palaka BK, Anithamma Kandukuri AK, Pamuru RR, Ampasala DR (2014) Response of antioxidative enzymes and lipoxygenase to drought stress in finger millet leaves (Eleusine coracana (L.) Gaertn). International Journal of Plant, Animal and Environmental Sciences 4, 644-653.
| Google Scholar |

Krishnamurthy L, Upadhyaya HD, Kashiwagi J, Purushothaman R, Dwivedi SL, Vadez V (2016a) Variation in drought-tolerance components and their interrelationships in the core collection of foxtail millet (Setaria italica) germplasm. Crop & Pasture Science 67(8), 834-846.
| Crossref | Google Scholar |

Krishnamurthy L, Upadhyaya HD, Kashiwagi J, Purushothaman R, Dwivedi SL, Vadez V (2016b) Variation in drought-tolerance components and their interrelationships in the minicore collection of finger millet germplasm. Crop Science 56(4), 1914-1926.
| Crossref | Google Scholar |

Kumawat KR, Sharma NK (2018) Effect of drought stress on plants growth. Popular Kheti 6(2), 239-241.
| Google Scholar |

Kusaka M, Lalusin AG, Fujimura T (2005) The maintenance of growth and turgor in pearl millet (Pennisetum glaucum [L.] Leeke) cultivars with different root structures and osmo-regulation under drought stress. Plant Science 168(1), 1-14.
| Crossref | Google Scholar |

Laxa M, Liebthal M, Telman W, Chibani K, Dietz K-J (2019) The role of the plant antioxidant system in drought tolerance. Antioxidants 8(4), 94.
| Crossref | Google Scholar |

Li J, Wang Y, Wang L, Zhu J, Deng J, Tang R, Chen G (2021) Integration of transcriptomic and proteomic analyses for finger millet [Eleusine coracana (L.) Gaertn.] in response to drought stress. PLoS ONE 16(2), e0247181.
| Crossref | Google Scholar |

Liu Z-J, Zhang X-L, Bai J-G, Suo B-X, Xu P-L, Wang L (2009) Exogenous paraquat changes antioxidant enzyme activities and lipid peroxidation in drought-stressed cucumber leaves. Scientia Horticulturae 121(2), 138-143.
| Crossref | Google Scholar |

Maharajan T, Antony Ceasar S, Ajeesh Krishna TP, Ignacimuthu S (2021) Finger millet [Eleusine coracana (L.) Gaertn]: an orphan crop with a potential to alleviate the calcium deficiency in the semi-arid tropics of Asia and Africa. Frontiers in Sustainable Food Systems 5, 684447.
| Crossref | Google Scholar |

Maharajan T, Krishna TPA, Rakkammal K, Ceasar SA, Ramesh M (2022) Application of CRISPR/Cas system in cereal improvement for biotic and abiotic stress tolerance. Planta 256(6), 106.
| Crossref | Google Scholar |

Mall TP, Tripathi SC (2016) Millets the nutrimental potent ethno-medicinal grasses: a review. World Journal of Pharmaceutical Research 5(2), 495-520.
| Google Scholar |

Maqsood M, Ali SNA (2007) Effects of drought on growth, development, radiation use efficiency and yield of finger millet (Eleucine coracana). Pakistan Journal of Botany 39(1), 123-134.
| Google Scholar |

Mazumder S, Bhattacharya D, Lahiri D, Nag M (2024) Milletomics: a metabolomics centered integrated omics approach toward genetic progression. Functional & Integrative Genomics 24(5), 149.
| Crossref | Google Scholar |

Merah O (2001) Potential importance of water status traits for durum wheat improvement under Mediterranean conditions. The Journal of Agricultural Science 137(2), 139-145.
| Crossref | Google Scholar |

Mickelbart MV, Hasegawa PM, Bailey-Serres J (2015) Genetic mechanisms of abiotic stress tolerance that translate to crop yield stability. Nature Reviews Genetics 16(4), 237-251.
| Crossref | Google Scholar | PubMed |

Mohanabharathi M, Sritharan N, Senthil A, Ravikesavan R (2019) Physiological studies for yield enhancement in finger millet under drought condition. Journal of Pharmacognosy and Phytochemistry 8(3), 3308-3312.
| Google Scholar |

Mude LN, Mondam M, Gujjula V, Jinka S, Pinjari OB, Yellodu Adi Reddy N, Patan SSVK (2020) Morpho-physiological and biochemical changes in finger millet [Eleusine coracana (L.) Gaertn.] under drought stress. Physiology and Molecular Biology of Plants 26(11), 2151-2171.
| Crossref | Google Scholar | PubMed |

Mukami A, Ngetich A, Mweu C, Oduor RO, Muthangya M, Mbinda WM (2019) Differential characterization of physiological and biochemical responses during drought stress in finger millet varieties. Physiology and Molecular Biology of Plants 25, 837-846.
| Crossref | Google Scholar | PubMed |

Mundada PS, Nikam TD, Anil Kumar S, Umdale SD, Ahire ML (2020) Morpho-physiological and biochemical responses of finger millet (Eleusine coracana (L.) Gaertn.) genotypes to PEG-induced osmotic stress. Biocatalysis and Agricultural Biotechnology 23, 101488.
| Crossref | Google Scholar |

Nagarjuna KN, Parvathi MS, Sajeevan RS, Pruthvi V, Mamrutha HM, Nataraja KN (2016) Full-length cloning and characterization of abiotic stress responsive CIPK31-like gene from finger millet, a drought-tolerant crop. Current Science 111, 890-894.
| Crossref | Google Scholar |

Nakanwagi MJ, Sseremba G, Kabod NP, Masanza M, Kizito EB (2020) Identification of growth stage-specific watering thresholds for drought screening in Solanum aethiopicum Shum. Scientific Reports 10(1), 862.
| Crossref | Google Scholar |

Noctor G, Foyer CH (1998) Ascorbate and glutathione: keeping active oxygen under control. Annual Review of Plant Physiology and Plant Molecular Biology 49(1), 249-279.
| Crossref | Google Scholar |

Noctor G, Veljovic-Jovanovic S, Foyer CH (2000) Peroxide processing in photosynthesis: antioxidant coupling and redox signalling. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences 355(1402), 1465-1475.
| Crossref | Google Scholar | PubMed |

Ozkur O, Ozdemir F, Bor M, Turkan I (2009) Physiochemical and antioxidant responses of the perennial xerophyte Capparis ovata Desf. to drought. Environmental and Experimental Botany 66(3), 487-492.
| Crossref | Google Scholar |

Panda D, Sailaja NH, Behera PK, Lenka K, Sharma SS, Lenka SK (2021a) Genetic diversity of under-utilized indigenous finger millet genotypes from Koraput, India for crop improvement. Journal of Plant Biochemistry and Biotechnology 30, 99-116.
| Crossref | Google Scholar |

Panda D, Rath C, Behera PK, Lenka SK (2021b) Physiological introspection of leaf photochemical activity and antioxidant metabolism in selected indigenous finger millet genotypes in relation to drought stress. Cereal Research Communications 49(4), 607-618.
| Crossref | Google Scholar |

Pandey V, Ansari MW, Tula S, Yadav S, Sahoo RK, Shukla N, Bains G, Badal S, Chandra S, Gaur AK, Kumar A, Shukla A, Kumar J, Tuteja N (2016) Dose-dependent response of Trichoderma harzianum in improving drought tolerance in rice genotypes. Planta 243, 1251-1264.
| Crossref | Google Scholar | PubMed |

Parvathi MS, Nataraja KN (2017) Discovery of stress responsive TATA-box binding protein associated Factor6 (TAF6) from finger millet (Eleusine coracana (L.) Gaertn). Journal of Plant Biology 60, 335-342.
| Crossref | Google Scholar |

Parvathi MS, Nataraja KN, Yashoda BK, Ramegowda HV, Mamrutha HM, Rama N (2013) Expression analysis of stress responsive pathway genes linked to drought hardiness in an adapted crop, finger millet (Eleusine coracana). Journal of Plant Biochemistry and Biotechnology 22, 193-201.
| Crossref | Google Scholar |

Parvathi MS, Nataraja KN, Reddy YAN, Naika MBN, Gowda MVC (2019) Transcriptome analysis of finger millet (Eleusine coracana (L.) Gaertn.) reveals unique drought responsive genes. Journal of Genetics 98(2), 46.
| Crossref | Google Scholar |

Pei Z-M, Murata Y, Benning G, Thomine S, Klüsener B, Allen GJ, Grill E, Schroeder JI (2000) Calcium channels activated by hydrogen peroxide mediate abscisic acid signalling in guard cells. Nature 406(6797), 731-734.
| Crossref | Google Scholar | PubMed |

Pérez-Arellano I, Carmona-Álvarez F, Martínez AI, Rodríguez-Díaz J, Cervera J (2010) Pyrroline-5-carboxylate synthase and proline biosynthesis: from osmotolerance to rare metabolic disease. Protein Science 19(3), 372-382.
| Crossref | Google Scholar | PubMed |

Pour-Aboughadareh A, Etminan A, Abdelrahman M, Siddique KHM, Tran L-SP (2020) Assessment of biochemical and physiological parameters of durum wheat genotypes at the seedling stage during polyethylene glycol-induced water stress. Plant Growth Regulation 92(1), 81-93.
| Crossref | Google Scholar |

Prabhakar M, Gopinath KA, Sai Sravan U, Srasvan Kumar G, Thirupathi M, Samba Siva G, Meghalakshmi G, Ravi Kumar N, Singh VK (2023) Potential for yield and soil fertility improvement with integration of organics in nutrient management for finger millet under rainfed Alfisols of Southern India. Frontiers in Nutrition 10, 1095449.
| Crossref | Google Scholar |

Pradhan A, Panda AK, Bhavani RV (2019) Finger millet in tribal farming systems contributes to increased availability of nutritious food at household level: insights from India. Agricultural Research 8(4), 540-547.
| Crossref | Google Scholar |

Quan L-J, Zhang B, Shi W-W, Li H-Y (2008) Hydrogen peroxide in plants: a versatile molecule of the reactive oxygen species network. Journal of Integrative Plant Biology 50(1), 2-18.
| Crossref | Google Scholar | PubMed |

Rakkammal K, Priya A, Pandian S, Maharajan T, Rathinapriya P, Satish L, Ceasar SA, Sohn S-I, Ramesh M (2022) Conventional and omics approaches for understanding the abiotic stress response in cereal crops – an updated overview. Plants 11(21), 2852.
| Crossref | Google Scholar |

Rakkammal K, Maharajan T, Shriram RN, Ram PSJ, Ceasar SA, Ramesh M (2023) Physiological, biochemical and molecular responses of finger millet (Eleusine coracana) genotypes exposed to short-term drought stress induced by PEG-6000. South African Journal of Botany 155, 45-59.
| Crossref | Google Scholar |

Ramakrishna C, Singh S, Raghavendrarao S, Padaria JC, Mohanty S, Sharma TR, Solanke AU (2018) The membrane tethered transcription factor EcbZIP17 from finger millet promotes plant growth and enhances tolerance to abiotic stresses. Scientific Reports 8(1), 2148.
| Crossref | Google Scholar |

Ramegowda V, Senthil-Kumar M, Nataraja KN, Reddy MK, Mysore KS, Udayakumar M (2012) Expression of a finger millet transcription factor, EcNAC1, in tobacco confers abiotic stress-tolerance. PLoS ONE 7(7), e40397.
| Crossref | Google Scholar |

Saleem S, Mushtaq NU, Shah WH, Rasool A, Hakeem KR, Rehman RU (2021) Morpho-physiological, biochemical and molecular adaptation of millets to abiotic stresses: a review. Phyton 90(5), 1363-1385.
| Crossref | Google Scholar |

Sandalio LM, Rodríguez-Serrano M, Romero-Puertas MC, del Río LA (2013) Role of peroxisomes as a source of reactive oxygen species (ROS) signaling molecules. In ‘Peroxisomes and their key role in cellular signaling and metabolism’. (Ed. L del Río) pp. 231–255. (Springer) 10.1007/978-94-007-6889-5_13

Sarker U, Oba S (2018) Drought stress effects on growth, ROS markers, compatible solutes, phenolics, flavonoids, and antioxidant activity in Amaranthus tricolor. Applied Biochemistry and Biotechnology 186, 999-1016.
| Crossref | Google Scholar | PubMed |

Sashidhar VR, Gurumurthy BR, Prasad TG, Udaya Kumar M, Seetharam A, Krishna Sastry KS (1986) Genotypic variation in carbon exchange rate, functional leaf area and productivity in finger millet (Eleusine coracana Gaertn.): an approach to identify desirable plant types for higher water use efficiency under rainfed conditions. Field Crops Research 13, 133-146.
| Crossref | Google Scholar |

Shao H, Wang H, Tang X (2015) NAC transcription factors in plant multiple abiotic stress responses: progress and prospects. Frontiers in Plant Science 6, 902.
| Crossref | Google Scholar |

Sharma P, Jha AB, Dubey RS, Pessarakli M (2012) Reactive oxygen species, oxidative damage, and antioxidative defense mechanism in plants under stressful conditions. Journal of Botany 2012(1), 217037.
| Crossref | Google Scholar |

Shin YK, Bhandari SR, Cho MC, Lee JG (2020) Evaluation of chlorophyll fluorescence parameters and proline content in tomato seedlings grown under different salt stress conditions. Horticulture, Environment, and Biotechnology 61, 433-443.
| Crossref | Google Scholar |

Shivhare R, Lata C (2019) Assessment of pearl millet genotypes for drought stress tolerance at early and late seedling stages. Acta Physiologiae Plantarum 41(3), 39.
| Crossref | Google Scholar |

Shrestha N, Hu H, Shrestha K, Doust AN (2023) Pearl millet response to drought: a review. Frontiers in Plant Science 14, 1059574.
| Crossref | Google Scholar |

Singh RK, Phanindra MLV, Singh VK, Sonam , Raghavendrarao S, Solanke AU, Kumar PA (2014) Isolation and characterization of drought responsive EcDehydrin7 gene from finger millet (Eleusine coracana (L.) Gaertn.). Indian Journal of Genetics and Plant Breeding 74(04), 456-462.
| Crossref | Google Scholar |

Singh RK, Singh VK, Raghavendrarao S, Phanindra MLV, Venkat Raman K, Solanke AU, Kumar PA, Sharma TR (2015) Expression of finger millet EcDehydrin7 in transgenic tobacco confers tolerance to drought stress. Applied Biochemistry and Biotechnology 177, 207-216.
| Crossref | Google Scholar | PubMed |

Sreeman SM, Vijayaraghavareddy P, Sreevathsa R, Rajendrareddy S, Arakesh S, Bharti P, Dharmappa P, Soolanayakanahally R (2018) Introgression of physiological traits for a comprehensive improvement of drought adaptation in crop plants. Frontiers in Chemistry 6, 92.
| Crossref | Google Scholar |

Srinivas V, Balasubramanian D (1995) Proline is a protein-compatible hydrotrope. Langmuir 11(7), 2830-2833.
| Crossref | Google Scholar |

Srinivasarao C, Venkateswarlu B, Kumar Singh A, Pandu Ranga Vittal K, Kundu S, Ravindra Chary G, Narayanaiyer Gajanan G, Kogganur Ramachandrappa B (2012) Critical carbon inputs to maintain soil organic carbon stocks under long-term finger-millet (Eleusine coracana [L.] Gaertn.) cropping on Alfisols in semiarid tropical India. Journal of Plant Nutrition and Soil Science 175(5), 681-688.
| Crossref | Google Scholar |

Sultan MARF, Hui L, Yang LJ, Xian ZH (2012) Assessment of drought tolerance of some Triticum L. species through physiological indices. Czech Journal of Genetics and Plant Breeding 48, 178-184.
| Crossref | Google Scholar |

Swapna S, Shylaraj KS (2017) Screening for osmotic stress responses in rice varieties under drought condition. Rice Science 24(5), 253-263.
| Crossref | Google Scholar |

Tadele Z (2016) ‘Drought adaptation in millets.’ (InTech)

Talwar HS, Kumar S, Madhusudhana R, Nanaiah GK, Ronanki S, Tonapi VA (2020) Variations in drought tolerance components and their association with yield components in finger millet (Eleusine coracana). Functional Plant Biology 47(7), 659-674.
| Crossref | Google Scholar | PubMed |

Tester M, Langridge P (2010) Breeding technologies to increase crop production in a changing world. Science 327(5967), 818-822.
| Crossref | Google Scholar | PubMed |

Turner NC (1996) Further progress in crop water relations. Advances in Agronomy 58, 293-338.
| Crossref | Google Scholar |

Tyagi J, Shrivastava N, Sharma AK, Varma A, Pudake R (2021) Effect of Rhizophagus intraradices on growth and physiological performance of Finger Millet (Eleusine coracana L.) under drought stress. Plant Science Today 8(4), 912-923.
| Crossref | Google Scholar |

Vadez V, Krishnamurthy L, Hash CT, Upadhyaya HD, Borrell AK (2011) Yield, transpiration efficiency, and water-use variations and their interrelationships in the sorghum reference collection. Crop & Pasture Science 62(8), 645-655.
| Crossref | Google Scholar |

Vetriventhan M, Upadhyaya HD, Dwivedi SL, Pattanashetti SK, Singh SK (2016) Finger and foxtail millets. In ‘Genetic and genomic resources for grain cereals improvement’. (Eds M Singh, HD Upadhyaya) pp. 291–319. (Elsevier)

Viljevac Vuletić M, Marček T, Španić V (2019) Photosynthetic and antioxidative strategies of flag leaf maturation and its impact to grain yield of two field-grown wheat varieties. Theoretical and Experimental Plant Physiology 31(3), 387-399.
| Crossref | Google Scholar |

Wang J, Li C, Li L, Reynolds M, Mao X, Jing R (2021) Exploitation of drought tolerance-related genes for crop improvement. International Journal of Molecular Sciences 22(19), 10265.
| Crossref | Google Scholar |

Yancey PH (2005) Organic osmolytes as compatible, metabolic and counteracting cytoprotectants in high osmolarity and other stresses. Journal of Experimental Biology 208(15), 2819-2830.
| Crossref | Google Scholar |

Yang J, Zhang J, Li C, Zhang Z, Ma F, Li M (2019) Response of sugar metabolism in apple leaves subjected to short-term drought stress. Plant Physiology and Biochemistry 141, 164-171.
| Crossref | Google Scholar | PubMed |

Yoon H-S, Lee H, Lee I-A, Kim K-Y, Jo J (2004) Molecular cloning of the monodehydroascorbate reductase gene from Brassica campestris and analysis of its mRNA level in response to oxidative stress. Biochimica et Biophysica Acta (BBA)-Bioenergetics 1658(3), 181-186.
| Crossref | Google Scholar |

Yuan S, Liu W-J, Zhang N-H, Wang M-B, Liang H-G, Lin H-H (2005) Effects of water stress on major photosystem II gene expression and protein metabolism in barley leaves. Physiologia Plantarum 125(4), 464-473.
| Crossref | Google Scholar |