A Systematic Review of Thousand-Grain Weight Genes in Rice and Foxtail Millet: Toward Functional Markers for Yield Improvement
DOI:
https://doi.org/10.29244/jtcs.13.03.664-680Keywords:
linkage marker, millet, PRISMA, SLR, SNPAbstract
Foxtail millet (Setaria italica (L.) P. Beauv.) is a climate-resilient crop with high potential for cultivation in low-input agricultural systems. However, its genetic improvement lags behind major crops such as rice. The objectives of this study were to identify agronomic traits associated with grain yield and to determine candidate genes for thousand-grain weight (TGW) in foxtail millet using a systematic literature review (SLR) combined with sequence based analysis. Rice was included in the SLR due to its close synteny with foxtail millet and its extensive genetic studies. In accordance with PRISMA 2020 guidelines, 116 articles were selected from the Scopus and PubMed databases. The results showed that TGW has a strong positive correlation with rice grain yield. Eight candidate genes (GS3, GW5, TGW6, GW2, GS5, GW8, GS2, and GW5L) controlling grain size and grain weight were identified based on their recurrence across studies and subsequently used to identify homologs in foxtail millet. The sequence of a foxtail millet homolog of TGW6, SiTGW6 (Seita.1G350400), was further analyzed to find potential variants and co-inherited genes. Sequence analysis identified SNPs in the SiTGW6 region, including a variant at 40,392,218 bp (MAF 0.17; moderate effect) that warrants further investigation for SNP based marker development. Analysis within the linkage disequilibrium (LD)-decay region identified Seita.1G350500.1, which contains BRCT, HAD-like, and FCP1 domains involved in transcriptional regulation. This study identifies SiTGW6 as a potential genetic target, with both SNP-based and linkage-marker development serving as a foundation for gene-based functional markers and prospective precision breeding in foxtail millet.
References
Akabane, T., Suzuki, N., Ikeda, K., Yonezawa, T., Nagatoishi, S., Matsumura, H., Yoshizawa, T., Tsuchiya, W., Kamino, S., Tsumoto, K., Ishimaru, K., Katoh, E., & Hirotsu, N. (2024). Thousand-grain weight 6, an IAA-glucose hydrolase, preferentially recognizes the indole ring structure. Scientific Reports, 14, 6778. https://doi.org/10.1038/s41598-024-57506-z
Ardie, S. W., Khumaida, N., Fauziah, N., & Yudiansyah. (2017). Biodiversity assessment of foxtail millet (Setaria italica L.) genotypes based on RAPD markers. Journal of Tropical Crop Science, 4(1), 21 26. https://doi.org/10.29244/jtcs.4.1.21-25
Ardie, S. W., Nugroho, R. B., Dirpan, A., & Anshori, M. F. (2025). Foxtail millet research in supporting climate change resilience efforts: Bibliometric analysis and focused literature review. Heliyon, 11(3), e42348. https://doi.org/10.1016/j.heliyon.2025.e42348
Arora, L., Aggarwal, R., Dhaliwal, I., Gupta, O. P., & Kaushik, P. (2023). Assessment of sensory and nutritional attributes of foxtail millet-based food products. Frontiers in Nutrition, 10, 1146545. https://doi.org/10.3389/fnut.2023.1146545
Ashfaq, L., Iqbal, S., Rasheed, S., Aasi, A. S., Niazi, A. K., & Khan, R. S. A. (2025). Bibliometric analysis of worldwide research trends on rice. Plant Bulletin, 4(1), 1–16. https://doi.org/10.55627/pbulletin.004.01.0905
Badro, H., Ndjiondjop, M. N., Furtado, A., & Henry, R. (2020). Sequence variants linked to key traits in interspecific crosses between African and Asian rice. Plants, 9(12), 1653. https://doi.org/10.3390/plants9121653
Baye, A., Berihun, B., Bantayehu, M., & Derebe, B. (2020). Genotypic and phenotypic correlation and path coefficient analysis for yield and yield-related traits in advanced bread wheat (Triticum aestivum L.) lines. Cogent Food & Agriculture, 6(1), 1752603. https://doi.org/10.1080/23311932.2020.1752603
Boell, S. K., & Cecez-Kecmanovic, D. (2015). On being ‘systematic’ in literature reviews in IS. Journal of Information Technology, 30(2), 161–173. https://doi.org/10.1057/jit.2014.26
Brignardello-Petersen, R., Santesso, N., & Guyatt, G. H. (2025). Systematic reviews of the literature: An introduction to current methods. American Journal of Epidemiology, 194(2), 536–542. https://doi.org/10.1093/aje/kwae232
Butarbutar, L. K. S., Syawaluddin, D. D., Suwarno, W. B., & Ardie, S. W. (2024). SiDREB2-based SNAP marker-assisted and multi-trait selection in the early generation of foxtail millet (Setaria italica L. Beauv.). Journal of Tropical Biodiversity and Biotechnology, 9(3), Article jtbb90407. https://doi.org/10.22146/jtbb.90407
Chibvongodze, M. K., Gasura, E., Kamutando, C. N., Mabasa, S., Makovere, B., Nyakudya, E., Nhamo, A., Mtambanengwe, F., & Mapfumo, P. (2025). Potential traits to aid selection for high grain yield based on trait associations and path analysis in sorghum. Journal of Underutilized Crops Research, 4(1), a23. https://doi.org/10.4102/jucr.v4i1.23
Cu, S. T., Warnock, N. I., Pasuquin, J., Dingkuhn, M., & Stangoulis, J. (2021). A high-resolution genome-wide association study of the grain ionome and agronomic traits in rice (Oryza sativa subsp. indica). Scientific Reports, 11, 19230. https://doi.org/10.1038/s41598-021-98573-w
Devos, K., Wang, Z., Beales, J., Sasaki, T., & Gale, M. D. (1998). Comparative genetic maps of foxtail millet (Setaria italica) and rice (Oryza sativa). Theoretical and Applied Genetics, 96(1), 63–68. https://doi.org/10.1007/s001220050709
Facchinello, P. H. K., Carvalho, I. R., Streck, E. A., Aguiar, G. A., Goveia, J., Feijo, M., Pereira, R. R., Fagundes, P. R. R., Loro, M. V., Maia, L. C., & Junior, A. M. M. (2023). Genetic trends and multivariate interrelationships for grain quality of irrigated rice genotypes. Agronomy Science and Biotechnology, 9, 1–16. https://doi.org/10.33158/ASB.r192. v9.2023
Falagas, M. E., Pitsouni, E. I., Malietzis, G. A., & Pappas, G. (2008). Comparison of PubMed, Scopus, Web of Science, and Google Scholar: Strengths and weaknesses. FASEB Journal, 22(2), 338 342. https://doi.org/10.1096/fj.07-9492LSF
Feng, Y., Lu, Q., Zhai, R., Zhang, M., Xu, Q., Yang, Y., Wang, S., Yuan, X., Yu, H., Wang, Y., & Wei, X. (2016). Genome-wide association mapping for grain shape traits in indica rice. Planta, 244(4), 819–830. https://doi.org/10.1007/s00425-016-2548-9
Gandhimeyyan, R. V., Chockalingam, V., Muthurajan, R., & Raman, R. (2023). Accelerating gene discovery in climate resilient and nutrient-rich major and minor millets through genome-wide association studies: Progress and prospects. Plant Breeding, 142(4), 389–406. https://doi.org/10.1111/pbr.13120
Gao, L., Zhu, Q., Li, H., Wang, S., Fan, J., Wang, T., Yang, L., Zhao, Y., Ma, Y., Chen, L., Li, X., Dong, S., Chu, X., Diao, X., Yuan, X., Wang, J., & Yang, G. (2025). Construction of a genetic linkage map and QTL mapping of the agronomic traits in foxtail millet (Setaria italica). BMC Genomics, 26, 152. https://doi.org/10.1186/s12864-024-11169-2
Gasparis, S., & Miłoszewski, M. M. (2023). Genetic basis of grain size and weight in rice, wheat, and barley. International Journal of Molecular Sciences, 24(23), 16921. https://doi.org/10.3390/ijms242316921
Gull, S., Haider, Z., Gu, H., Khan, R. A. R., Miao, J., Wenchen, T., Uddin, S., Ahmad, I., & Liang, G. (2019). InDel marker-based estimation of multi-gene allele contribution and genetic variations for grain size and weight in rice (Oryza sativa L.). International Journal of Molecular Sciences, 20(19), 4824. https://doi.org/10.3390/ijms20194824
Han, Y., Luo, D., Usman, B., Nawaz, G., Zhao, N., Liu, F., & Li, R. (2018). Development of high-yielding glutinous cytoplasmic male sterile rice (Oryza sativa L.) lines through CRISPR/Cas9-based mutagenesis of Wx and TGW6, and proteomic analysis of anthers. Agronomy, 8(12), 290. https://doi.org/10.3390/agronomy8120290
Hou, X., Chen, M., Chen, Y., Hou, X., Jia, Z., Yang, X., Zhang, J., Liu, Y., & Ye, N. (2022). Identifying QTLs for grain size in a colossal-grain rice (Oryza sativa L.) line and analyzing the additive effects of QTLs. International Journal of Molecular Sciences, 23(7), 3526. https://doi.org/10.3390/ijms23073526
Jaiswal, V., Gupta, S., Gahlaut, V., Muthamilarasan, M., Bandyopadhyay, T., Ramchiary, N., & Prasad, M. (2019). Genome-wide association study of major agronomic traits in foxtail millet (Setaria italica L.) using ddRAD sequencing. Scientific Reports, 9, 5020. https://doi.org/10.1038/s41598-019-41602-6
Jannah, R. M., Ratnawati, S., Suwarno, W. B., & Ardie, S. W. (2024). Digital phenotyping for robust seeds variability assessment in Setaria italica (L.) P. Beauv. Journal of Seed Science, 46, e281586. https://doi.org/10.1590/2317-1545v46281586
Kalsi, R., & Bhasin, J. K. (2023). Nutritional exploration of foxtail millet (Setaria italica) in addressing food security and its utilization trends in food systems. eFood, 4(5), e111. https://doi.org/10.1002/efd2.111
Karamang, S., Widaryanti, A. E., & Aini, N. (2023). Yield quality of Setaria italica accessions originated from Numfor Island, Papua, Indonesia. Biodiversitas, 24(3), 1878–1885. https://doi.org/10.13057/biodiv/d240364
Kasule, F., Diack, O., Mbaye, M., Kakeeto, R., & Econopouly, B. F. (2024). Genomic resources, opportunities, and prospects for accelerated improvement of millets. Theoretical and Applied Genetics, 137, 273. https://doi.org/10.1007/s00122-024-04777-9
Kumar, G. P., Sunil, N., Srinivasa Chary, D., & Sekhar, J. C. (2024). Character association and path coefficient analysis to determine interrelationships among grain yield and its components in maize genotypes (Zea mays L.). Journal of Scientific Research and Reports, 30(3), 275–283. https://doi.org/10.9734/jsrr/2024/v30i31878
Lata, C., Bhutty, S., Bahadur, R. P., Majee, M., & Prasad, M. (2011). Association of an SNP in a novel DREB2-like gene SiDREB2 with stress tolerance in foxtail millet [Setaria italica (L.)]. Journal of Experimental Botany, 62(10), 3387–3401. https://doi.org/10.1093/jxb/err016
Liang, H., He, Q., Zhang, H., Zhi, H., Tang, S., Wang, H., Meng, Q., Jia, G., Chang, J., & Diao, X. (2023). Identification and haplotype analysis of SiCHLI: A gene for yellow-green seedlings as a morphological marker to accelerate foxtail millet (Setaria italica) hybrid breeding. Theoretical and Applied Genetics, 136(1), 24. https://doi.org/10.1007/s00122-023-04309-x
Liang, S., Sun, C., Ren, G., Zhao, X., Zhou, M., McNeil, D., & Ye, G. (2016). Usefulness of the cloned and fine-mapped genes/QTL for grain yield and related traits in indica rice breeding for irrigated ecosystems. Field Crops Research, 187, 58–73. https://doi.org/10.1016/j.fcr.2015.11.014
Liang, Y., & Han, Y. (2024). The pan-genome offers opportunities to revitalize the ancient crop foxtail millet. Plant Communications, 5. https://doi.org/10.1016/j.xplc.2023.100735
Liu, G., Qiu, D., Lu, Y., Wu, Y., Han, X., Jiao, Y., Wang, T., Yang, J., You, A., Chen, J., & Zhang, Z. (2023). Identification of superior haplotypes and haplotype combinations for grain size- and weight-related genes for breeding applications in rice (Oryza sativa L.). Genes, 14(12), 2201. https://doi.org/10.3390/genes14122201
Liu, T., He, J., Dong, K., Wang, X., Wang, W., Yang, P., Ren, R., Zhang, L., Zhang, Z., & Yang, T. (2020). QTL mapping of yield component traits on bin map generated from resequencing a RIL population of foxtail millet (Setaria italica). BMC Genomics, 21(1), 141. https://doi.org/10.1186/s12864-020-6553-9
Liu, X., Yang, Y., Hou, S., Men, Y., & Han, Y. (2022). The integration of genome-wide association study and homology analysis to explore the genomic regions and candidate genes for panicle-related traits in foxtail millet. International Journal of Molecular Sciences, 23(23), 14735. https://doi.org/10.3390/ijms232314735
Luthfiani, M. V., Pahlevi, M. R., Purwoko, B. S., & Ardie, S. W. (2025). Insights into panicle trait variation and conservation of the DUF-640 gene in Indonesian foxtail millets (Setaria italica). Czech Journal of Genetics and Plant Breeding, 61(4), 235–246. https://doi.org/10.17221/86/2025-CJGPB
Manners, R., & van Etten, J. (2018). Are agricultural researchers working on the right crops to enable food and nutrition security under future climates? Global Environmental Change, 53, 182–194. https://doi.org/10.1016/j.gloenvcha.2018.09.010
Muzzayyanah, P. N., Suwarno, W. B., & Ardie, S. W. (2024). Gene action and heritability estimates in F₂ populations of foxtail millet (Setaria italica L.). SABRAO Journal of Breeding and Genetics, 56(1), 65–75. https://doi.org/10.54910/sabrao2024.56.1.6
Nagaraju, B., Basavaraj, K., Gireesh, C., & Sasipriya, S. (2023). Variability parameters, correlation studies and path analysis of yield and yield-related traits in rice (Oryza sativa L.): A comprehensive review. International Journal of Environment and Climate Change, 13(11), 2015–2022. https://doi.org/10.9734/ijecc/2023/v13i113360
Niu, Y., Chen, T., Wang, C., Chen, K., Shen, C., Chen, H., Zhu, S., Wu, Z., Zheng, T., Zhang, F., & Xu, J. (2021). Identification and allele mining of new candidate genes underlying rice grain weight and grain shape by genome-wide association study. BMC Genomics, 22, 602. https://doi.org/10.1186/s12864-021-07901-x
Ouzzani, M., Hammady, H., Fedorowicz, Z., & Elmagarmid, A. (2016). Rayyan—A web and mobile app for systematic reviews. Systematic Reviews, 5(1), 210. https://doi.org/10.1186/s13643-016-0384-4
Page, M. J., McKenzie, J. E., Bossuyt, P. M., Boutron, I., Hoffmann, T. C., Mulrow, C. D., Shamseer, L., Tetzlaff, J. M., Akl, E. A., Brennan, S. E., Chou, R., Glanville, J., Grimshaw, J. M., Hróbjartsson, A., Lalu, M. M., Li, T., Loder, E. W., Mayo Wilson, E., McDonald, S., McGuinness, L. A., et al. (2021). The PRISMA 2020 statement: An updated guideline for reporting systematic reviews. BMJ, 372, n71. https://doi.org/10.1136/bmj.n71
Pareek, S., Pheirim, R., Chetariya, C. P., & Soharu, A. (2026). A comprehensive review of approaches for genetic improvement in foxtail millet (Setaria italica L.). Genetic Resources, 7(13), 169–180. https://doi.org/10.46265/genresj.OVEO3504
Ponce, K., Zhang, Y., Guo, L., Leng, Y., & Ye, G. (2020). Genome-wide association study of grain-size traits in the Indica rice multiparent advanced generation intercross (MAGIC) population. Frontiers in Plant Science, 11, 395. https://doi.org/10.3389/fpls.2020.00395
Rachmandika, F. C. A., Satrio, R. D., Hamim, H., & Miftahudin, M. (2025). SNP-based molecular characterization of TAD1, GS3, and ABA8ox2 genes in Indonesian rice for early marker discovery. IOP Conference Series: Earth and Environmental Science, 1550(1), 012047. https://doi.org/10.1088/1755-1315/1550/1/012047
Ramlah, Pabendon, M. B., & Daryono, B. S. (2020). Local food diversification of foxtail millet (Setaria italica) cultivars in West Sulawesi, Indonesia: A case study of diversity and local culture. Biodiversitas, 21(1), 67–73. https://doi.org/10.13057/biodiv/d210110
Ratnawati, S., Jannah, R. M., Dewi, Y. I., Rizqullah, R., Suwarno, W. B., & Ardie, S. W. (2024). The genetic variability of Indonesian local foxtail millet accession based on agro-morphological traits and early salinity tolerance evaluation utilizing SiDREB2-based SNAP marker. Hayati Journal of Biosciences, 31(1), 82–93. https://doi.org/10.4308/hjb.31.1.82-93
Salgotra, R. K., & Stewart, C. N., Jr. (2020). Functional markers for precision plant breeding. International Journal of Molecular Sciences, 21(13), 4792. https://doi.org/10.3390/ijms21134792
Semeskandi, M. N., Mazloom, P., Arabzadeh, B., Moghadam, M. N., & Ahmadi, T. (2024). Application of correlation coefficients and principal components analysis in the stability of quantitative and qualitative traits in rice improvement cultivation. Brazilian Journal of Biology, 84, e268981. https://doi.org/10.1590/1519-6984.268981
Sintia, M., Suwarno, W. B., & Ardie, S. W. (2023). Genetic variability of F₂ foxtail millet population derived from ICERI 5 and Botok 10 cross. Biodiversitas, 24(6), 3559–3567. https://doi.org/10.13057/biodiv/d240655
Suroya, L. F., Wirnas, D., Trikoesoemaningtyas, & Reflinur. (2023). Identification of waxy genotype in sorghum genetic resources using waxy gene-based markers and iodine staining methods. Australian Journal of Crop Science, 17(2), 190–197. https://doi.org/10.21475/ajcs.23.17.02.p3784
Suwarno, W. B., Pixley, K. V., Palacios-Rojas, N., Kaeppler, S. M., & Babu, R. (2015). Genome-wide association analysis reveals new targets for carotenoid biofortification in maize. Theoretical and Applied Genetics, 128(5), 851–864. https://doi.org/10.1007/s00122-015-2475-3
Usman, B., Zhao, N., Nawaz, G., Qin, B., Liu, F., Liu, Y., & Li, R. (2021). CRISPR/Cas9 guided mutagenesis of grain size 3 confers increased rice (Oryza sativa L.) grain length by regulating cysteine proteinase inhibitor and ubiquitin-related proteins. International Journal of Molecular Sciences, 22(6), 3225. https://doi.org/10.3390/ijms22063225
Wang, Y., Pang, Y., Chen, K., Zhai, L., Shen, C., Wang, S., & Xu, J. (2020). Genetic bases of source-, sink-, and yield-related traits revealed by a genome-wide association study in Xian rice. The Crop Journal, 8(1), 119–131. https://doi.org/10.1016/j.cj.2019.05.001
Wang, Y., Zheng, Y., Cai, Q., Liao, C., Mao, X., Xie, H., Zhu, Y., Lian, L., Luo, X., Xie, H., & Zhang, J. (2016). Population structure and association analysis of yield and grain quality traits in hybrid rice primal parental lines. Euphytica, 212, 261–273. https://doi.org/10.1007/s10681-016-1766-3
Wang, Z., Du, X., Wang, J., Yang, H., Wang, X., Guo, E., Wang, Y., Yuan, F., Tian, G., Liu, X., Wang, Q., Li, H., Zhang, L., & Peng, S. (2017). Molecular cloning, expression analysis, and development of functional markers for the SiARGOS1 gene in foxtail millet. Scientia Agricultura Sinica, 50(22), 4266–4276. https://doi.org/10.3864/j.issn.0578-1752.2017.22.003
Widyawan, M. H., Khumaida, N., Kitashiba, H., Nishio, T., & Ardie, S. W. (2018). Optimization of dot-blot SNP analysis for detecting drought- or salinity-stress associated markers in foxtail millet (Setaria italica L.). SABRAO Journal of Breeding and Genetics, 50(1), 72–84.
Xu, F., Sun, X., Chen, Y., Huang, Y., Tong, C., & Bao, J. (2015). Rapid identification of major QTLs associated with rice grain weight and their utilization. PLOS ONE, 10(3), e0122206. https://doi.org/10.1371/journal.pone.0122206
Xue, X., Yu, Z., Mu, A., Yang, F., Liu, D., Zhang, S., Zhang, J., Cheng, Y., Zhao, Y., Zhang, Y., & Wang, X. (2025). Identification and evaluation of phenotypic characters and genetic diversity analysis of 1,558 foxtail millet germplasm resources for conservation and breeding. Frontiers in Plant Science, 16, 1624252. https://doi.org/10.3389/fpls.2025.1624252
Yang, L., Wang, Y., Jahan, N., Hu, H., Chen, P., Shang, L., Lin, H., Dong, G., Hu, J., Gao, Z., Qian, Q., Zhang, Y., & Guo, L. (2019). Genome-wide association analysis and allelic mining of grain shape-related traits in rice. Rice Science, 26(6), 384-392. https://doi.org/10.1016/j.rsci.2018.09.002
Yang, T., Dong, J., Xiong, X., Zhang, L., Wang, J., Hu, H., Zhou, L., Yang, W., Ma, Y., Fu, H., Chen, J., Li, W., Nie, S., Liu, Z., Liu, B., Wang, F., Zhao, J., & Zhang, S. (2024). A novel function of GW5 on controlling early growth vigor and its haplotype effect on shoot dry weight and grain size in rice (Oryza sativa L.). Rice, 17, 49. https://doi.org/10.1186/s12284-024-00728-6
Yang, W., Zhang, F., Zafar, S., Wang, J., Lu, H., Naveed, S., Lou, J., & Xu, J. (2021). Genetic dissection of heterosis of indica japonica by introgression line, recombinant inbred line and their testcross populations. Scientific Reports, 11, 10265. https://doi.org/10.1038/s41598-021-89691-6
Zhang, G., Liu, X., Quan, Z., Cheng, S., Xu, X., Pan, S., Xie, M., Zeng, P., Yue, Z., Wang, W., Tao, Y., Bian, C., Han, C., Xia, Q., Peng, X., Cao, R., Yang, X., Zhan, D., Hu, J., Zhang, Y., et al. (2012). The genome sequence of foxtail millet (Setaria italica) provides insights into grass evolution and biofuel potential. Nature Biotechnology, 30(6), 549–554. https://doi.org/10.1038/nbt.2195
Zhang, H., Tang, S., Wang, H., Wang, Y., Zhi, H., Liu, B., Zhang, R., Ma, Q., Jia, G., Feng, B., & Diao, X. (2024). Genetic diversity in grain yield traits and identification of a grain weight gene, SiTGW6, in foxtail millet. Theoretical and Applied Genetics, 137, 84. https://doi.org/10.1007/s00122-024-04586-0
Zhang, W., Wang, B., Liu, B., Chen, Z., Lu, G., Ge, Y., & Bai, C. (2022). Trait selection for yield improvement in foxtail millet (Setaria italica Beauv.) under climate change in the North China Plain. Agronomy, 12(7), 1500. https://doi.org/10.3390/agronomy12071500
Zhao, Q., Shi, X. S., Wang, T., Chen, Y., Yang, R., Mi, J., Zhang, Y. W., & Zhang, Y. M. (2023). Identification of QTNs, QTN-by environment interactions, and candidate genes for grain size traits in main-crop and ratoon rice. Frontiers in Plant Science, 14, 1119218. https://doi.org/10.3389/fpls.2023.1119218
Zhong, H., Liu, C., Kong, W., Zhang, Y., Zhao, G., Sun, T., & Li, Y. (2020). Effect of multi allele combination on rice grain size based on prediction of regression equation model. Molecular Genetics and Genomics, 295(2), 465–474. https://doi.org/10.1007/s00438-019-01627-y
Zhong, H., Liu, S., Sun, T., Kong, W., Deng, X., Peng, Z., & Li, Y. (2021). Multi-locus genome-wide association studies for five yield-related traits in rice. BMC Plant Biology, 21, 364. https://doi.org/10.1186/s12870-021-03146-8
Downloads
Published
How to Cite
Issue
Section
License
All publications by Journal of Tropical Crop Science is licensed under a Creative Commons Attribution-ShareAlike 4.0 International License.



