Ghosh, S., Suri, D. & Uauy, R. Assessment of protein adequacy in developing countries: quality matters. Br. J. Nutr. 108, S77–S87 (2012).

Article 
CAS 
PubMed 

Google Scholar
 

Joint FAO/WHO/UNU Expert Consultation. Protein and Amino Acid Requirements in Human Nutrition: Report of a Joint FAO/WHO/UNU Expert Consultation (World Health Organization, 2007).

Galili, G., Amir, R. & Fernie, A. R. The regulation of essential amino acid synthesis and accumulation in plants. Annu. Rev. Plant Biol. 67, 153–178 (2016).

Article 
CAS 
PubMed 

Google Scholar
 

Papier, K. et al. Meat consumption and risk of 25 common conditions: outcome-wide analyses in 475,000 men and women in the UK Biobank study. BMC Med. 19, 53 (2021).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Simojoki, M. et al. The impacts of partial replacement of red and processed meat with legumes or cereals on protein and amino acid intakes: a modelling study in the Finnish adult population. Ann. Med. 55, 2281661 (2023).

Article 
PubMed 
PubMed Central 

Google Scholar
 

Rockström, J. et al. The EAT–Lancet Commission on healthy, sustainable, and just food systems. Lancet 406, 1625–1700 (2025).

Article 
PubMed 

Google Scholar
 

World Population Projected to Reach 9.8 Billion in 2050, and 11.2 Billion in 2100 (United Nations, 2017).

Yu, J. et al. White rice, brown rice and the risk of type 2 diabetes: a systematic review and meta-analysis. BMJ Open 12, e065426 (2022).

Article 
PubMed 
PubMed Central 

Google Scholar
 

Anjana, R. M. et al. Dietary profiles and associated metabolic risk factors in India from the ICMR-INDIAB survey-21. Nat. Med. 31, 3813–3824 (2025).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Tiozon, R. N. et al. Machine learning technique unraveled subspecies-specific ionomic variation with the preferential mineral enrichment in rice. Cereal Chem. 101, 367–381 (2024).

Article 
CAS 

Google Scholar
 

Kaur, R. et al. Protein profiling in a set of wild rice species and rice cultivars: a stepping stone to protein quality improvement. Cereal Res. Commun. 51, 163–177 (2023).

Article 
CAS 

Google Scholar
 

Long, X., Liu, Q., Chan, M., Wang, Q. & Sun, S. S. Metabolic engineering and profiling of rice with increased lysine. Plant Biotechnol. J. 11, 490–501 (2013).

Article 
CAS 
PubMed 

Google Scholar
 

Mosleth, E. F. et al. Genetic variation and heritability of grain protein deviation in European wheat genotypes. Field Crops Res. 255, 107896 (2020).

Article 
PubMed 
PubMed Central 

Google Scholar
 

Zhou, B., Serret, M. D., Pie, J. B., Shah, S. S. & Li, Z. Relative contribution of nitrogen absorption, remobilization, and partitioning to the ear during grain filling in Chinese winter wheat. Front. Plant Sci. 9, 1351 (2018).

Article 
PubMed 
PubMed Central 

Google Scholar
 

Padhan, B. K., Sathee, L., Kumar, S., Chinnusamy, V. & Kumar, A. Variation in nitrogen partitioning and reproductive stage nitrogen remobilization determines nitrogen grain production efficiency (NUEg) in diverse rice genotypes under varying nitrogen supply. Front. Plant Sci. 14, 1093581 (2023).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Peng, B. et al. OsAAP6 functions as an important regulator of grain protein content and nutritional quality in rice. Nat. Commun. 5, 4847 (2014.

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Ji, Y., Huang, W., Wu, B., Fang, Z. & Wang, X. The amino acid transporter AAP1 mediates growth and grain yield by regulating neutral amino acid uptake and reallocation in Oryza sativa. J. Exp. Bot. 71, 4763–4777 (2020).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Wang, T. et al. Amino acid permease 6 regulates grain protein content in maize. Crop J. 10, 1536–1544 (2022).

Article 
CAS 

Google Scholar
 

Peng, B. et al. OsAAP8 mutation leads to significant improvement in the nutritional quality and appearance of rice grains. Mol. Breed. 44, 34 (2024).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Wan, Y. F. et al. Wheat amino acid transporters highly expressed in grain cells regulate amino acid accumulation in grain. PLoS ONE 16, e0246763 (2021).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Peng, B. et al. OsNAC74 modulates rice growth and salt tolerance via hormone signaling pathways. Sci. Rep. 15, 45350 (2025).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Yang, N. et al. Two teosintes made modern maize. Science 382, eadg8940 (2023).

Article 
CAS 
PubMed 

Google Scholar
 

Flint-Garcia, S. A., Bodnar, A. L. & Scott, M. P. Wide variability in kernel composition, seed characteristics, and zein profiles among diverse maize inbreds, landraces, and teosinte. Theor. Appl. Gen. 119, 1129–1142 (2009).

Article 

Google Scholar
 

Huang, Y. et al. THP9 enhances seed protein content and nitrogen-use efficiency in maize. Nature 612, 292–300 (2022).

Article 
CAS 
PubMed 

Google Scholar
 

Lee, S. et al. OsASN1 overexpression in rice increases grain protein content and yield under nitrogen-limiting conditions. Plant Cell Physiol. 61, 1309–1320 (2020).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Hu, B. et al. Variation in NRT1.1B contributes to nitrate-use divergence between rice subspecies. Nat. Genet. 47, 834–838 (2015).

Article 
CAS 
PubMed 

Google Scholar
 

Cao, H. R. et al. ZmNRT1.1B (ZmNPF6.6) determines nitrogen use efficiency via regulation of nitrate transport and signalling in maize. Plant Biotechnol. J. 22, 316–329 (2024).

Article 
CAS 
PubMed 

Google Scholar
 

Wu, J. et al. Nature variations of OsNLP4 responsible for nitrogen use efficiency divergence in the two rice subspecies. Nat. Commun. 16, 7681 (2025).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Zhang, Z. S. et al. Rice NIN-LIKE PROTEIN 3 modulates nitrogen use efficiency and grain yield under nitrate-sufficient conditions. Plant Cell Environ. 45, 1520–1536 (2022).

Article 
CAS 
PubMed 

Google Scholar
 

Ding, Z., Wang, C., Chen, S. & Yu, S. Diversity and selective sweep in the OsAMT1;1 genomic region of rice. BMC Evol. Biol. 11, 61 (2011).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Xu, G., Fan, X. & Miller, A. J. Plant nitrogen assimilation and use efficiency. Annu. Rev. Plant Biol. 63, 153–182 (2012).

Article 
CAS 
PubMed 

Google Scholar
 

Lundstrom, M., Leino, M. W. & Hagenblad, J. Evolutionary history of the NAM-B1 gene in wild and domesticated tetraploid wheat. BMC Genet. 18, 118 (2017).

Article 
PubMed 
PubMed Central 

Google Scholar
 

Cha, J.-K. et al. Genotyping the high protein content gene NAM-B1 in wheat (Triticum aestivum L.) and the development of a KASP marker to identify a functional haplotype. Agronomy 13, 1977 (2023).

Article 
CAS 

Google Scholar
 

Roncallo, P. F. et al. Allelic variation at glutenin loci (Glu-1, Glu-2 and Glu-3) in a worldwide durum wheat collection and its effect on quality attributes. Foods 10, 2845 (2021).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Shrestha, V. et al. Multiomics approach reveals a role of translational machinery in shaping maize kernel amino acid composition. Plant Physiol. 188, 111–133 (2022).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Yang, T., Wu, X., Wang, W. & Wu, Y. Regulation of seed storage protein synthesis in monocot and dicot plants: a comparative review. Mol. Plant 16, 145–167 (2023).

Article 
CAS 
PubMed 

Google Scholar
 

Sreenivasulu, N. & Wobus, U. Seed-development programs: a systems biology-based comparison between dicots and monocots. Annu. Rev. Plant Biol. 64, 189–217 (2013).

Article 
CAS 
PubMed 

Google Scholar
 

Chen, E. R. et al. The transcription factors ZmNAC128 and ZmNAC130 coordinate with Opaque2 to promote endosperm filling in maize. Plant Cell 35, 4066–4090 (2023).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Peng, D. et al. Central roles of ZmNAC128 and ZmNAC130 in nutrient uptake and storage during maize grain filling. Genes 15, 663 (2024).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Luo, G. B. et al. Genome-wide identification of seed storage protein gene regulators in wheat through coexpression analysis. Plant J. 108, 1704–1720 (2021).

Article 
CAS 
PubMed 

Google Scholar
 

Wang, X. et al. Wheat NAC-A18 regulates grain starch and storage proteins synthesis and affects grain weight. Theor. Appl. Genet. 136, 123 (2023).

Article 
CAS 
PubMed 

Google Scholar
 

Xie, L. et al. Efficient proteome-wide identification of transcription factors targeting Glu-1: a case study for functional validation of TaB3-2A1 in wheat. Plant Biotechnol. J. 21, 1952–1965 (2023).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Guo, D. et al. Over-expressing TaSPA-B reduces prolamin and starch accumulation in wheat (Triticum aestivum L.) grains. Int. J. Mol. Sci. 21, 3257 (2020).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Sreenivasulu, N., Zhang, C., Tiozon, R. N. Jr. & Liu, Q. Post-genomics revolution in the design of premium quality rice in a high-yielding background to meet consumer demands in the 21st century. Plant Commun. 3, 100271 (2022).

Article 
CAS 
PubMed 

Google Scholar
 

Shi, Y. et al. Natural variations of OsAUX5, a target gene of OsWRKY78, control the neutral essential amino acid content in rice grains. Mol. Plant 16, 322–336 (2023).

Article 
CAS 
PubMed 

Google Scholar
 

Zhang, C. Q. et al. The WRKY transcription factor OsWRKY78 regulates stem elongation and seed development in rice. Planta 234, 541–554 (2011).

Article 
CAS 
PubMed 

Google Scholar
 

Atlin, G. et al. Quality protein maize: progress and prospects. Plant Breed. Rev. 34, 83–130 (2011).

CAS 

Google Scholar
 

Salazar-Salas, N. Y. et al. Biochemical characterization of QTLs associated with endosperm modification in quality protein maize. J. Cereal Sci. 60, 255–263 (2014).

Article 
CAS 

Google Scholar
 

Maqbool, M. A., Issa, A. B. & Khokhar, E. S. Quality protein maize (QPM): importance, genetics, timeline of different events, breeding strategies and varietal adoption. Plant Breed. 140, 375–399 (2021).

Article 
CAS 

Google Scholar
 

Badoni, S. et al. Multiomics of a rice population identifies genes and genomic regions that bestow low glycemic index and high protein content. Proc. Natl Acad. Sci. USA 121, e2410598121 (2024).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Li, Y. F. et al. Coordinated improvement of maize grain yield and protein quality by the ZmMADS8–ZmMADS47–O2 module and a G protein gamma subunit. Crop J. 13, 805–817 (2025).

Article 

Google Scholar
 

Lu, X. et al. Natural variations in the promoter of ZmDeSI2 encoding a deSUMOylating isopeptidase controls kernel methionine content in maize. Mol. Plant 18, 872–891 (2025).

Article 
CAS 
PubMed 

Google Scholar
 

Hogy, P. et al. Effects of elevated CO2 on grain yield and quality of wheat: results from a 3-year free-air CO2 enrichment experiment. Plant Biol. 11, 60–69 (2009).

Article 
PubMed 

Google Scholar
 

Wang, J. et al. Changes in grain protein and amino acids composition of wheat and rice under short-term increased [CO2] and temperature of canopy air in a paddy from East China. New Phytol. 222, 726–734 (2019).

Article 
CAS 
PubMed 

Google Scholar
 

Saha, S., Chakraborty, D., Sehgal, V. K. & Pal, M. Potential impact of rising atmospheric CO2 on quality of grains in chickpea (Cicer arietinum L.). Food Chem. 187, 431–436 (2015).

Article 
CAS 
PubMed 

Google Scholar
 

Li, W. et al. A natural gene on–off system confers field thermotolerance for grain quality and yield in rice. Cell 188, 4170–4172 (2025).

Article 
CAS 
PubMed 

Google Scholar
 

Tessari, P., Lante, A. & Mosca, G. Essential amino acids: master regulators of nutrition and environmental footprint. Sci. Rep. 6, 26074 (2016).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Trumbo, P., Schlicker, S., Yates, A. A. & Poos, M. Dietary reference intakes for energy, carbohydrate, fiber, fat, fatty acids, cholesterol, protein, and amino acids. J. Am. Diet Assoc. 102, 1621–1630 (2002).

Article 
PubMed 

Google Scholar
 

Poore, J. & Nemecek, T. Reducing food’s environmental impacts through producers and consumers. Science 360, 987–992 (2018).

Article 
CAS 
PubMed 

Google Scholar
 

Mekonnen, M. M. & Hoekstra, A. Y. The green, blue and grey water footprint of crops and derived crop products. Hydrol. Earth Syst. Sci. 15, 1577–1600 (2011).

Article 

Google Scholar
 

Mekonnen, M. M. & Hoekstra, A. Y. The Green, Blue and Grey Water Footprint of Farm Animals and Animal Products. Value of Water Research Report Series No. 48 (UNESCO-IHE, 2010).

FAOSTAT (FAO, 2026); https://www.fao.org/faostat

Production, Supply and Distribution (USDA Foreign Agriculture Service, 2026); https://apps.fas.usda.gov/psdonline

Bhavadharini, B. et al. White rice intake and incident diabetes: a study of 132,373 participants in 21 countries. Diabetes Care 43, 2643–2650 (2020).

Article 
PubMed 
PubMed Central 

Google Scholar
 

Ekpa, O., Palacios-Rojas, N., Kruseman, G., Fogliano, V. & Linnemann, A. R. Sub-Saharan African maize-based foods-processing practices, challenges and opportunities. Food Rev. Int. 35, 609–639 (2019).

Article 

Google Scholar
 

Erenstein, O. et al. in Wheat Improvement: Food Security in a Changing Climate (eds Reynolds, M. P., Sadras, V. O. & Lobell, D. B.) 47–66 (Springer International, 2022).

Uauy, C., Brevis, J. C. & Dubcovsky, J. The high grain protein content gene Gpc-B1 accelerates senescence and has pleiotropic effects on protein content in wheat. J. Exp. Bot. 57, 2785–2794 (2006).

Article 
CAS 
PubMed 

Google Scholar
 

Global Burden of Disease Study 2021 (GBD 2021) Data Resources (IHME, 2021).

EDGAR—Emissions Database for Global Atmospheric Research (European Commission, 2024).