Please wait a minute...
Journal of Integrative Agriculture
Advanced Online Publication | Current Issue | Archive | Adv Search
Advances in rice synthetic biology: toward a better staple crop and beyond

Chenchen Zhang, Yan Wang, Lu Chen, Xixi Wang, Sheng Teng#

Key Laboratory of Microbiological Metrology, Measurement & Bio-product Quality Security, State Administration for Market Regulation, College of Life Sciences, China Jiliang University, Hangzhou 310018, China

 Highlights 

• Expansion of synthetic biology into rice reflects its growing role in addressing global challenges in agriculture and human health.

• Substantial advancements in rice synthetic biology have enhanced yield, nutritional quality, stress resistance, and resource-use efficiency.

• Synthetic production of high-value micronutrients, pharmaceutical compounds, and therapeutic proteins or peptides in the endosperm positions rice for roles extending beyond food security.

Download:  PDF in ScienceDirect  
Export:  BibTeX | EndNote (RIS)      
摘要  

合成生物学是一门运用工程学原理设计并构建新型生物系统或生命体的交叉学科。合成生物学最初以微生物系统为主要研究对象,目前已拓展至植物体系。植物合成生物学为应对农业与人口健康领域的全球性挑战提供了富有前景的解决方案。水稻作为全球主要粮食作物及植物科学研究的模式物种,已成为该领域的关键研究平台。近年来,研究人员在基于多重基因组编辑的基因组工程、新一代杂交稻的合成、无融合生殖的诱导、光合与固氮途径的重构,以及微量营养素、药物分子、治疗性蛋白与多肽的生物合成等方面取得了重要进展。本文系统综述了水稻合成生物学领域的最新研究成果,总结了当前发展态势,并对未来研究方向进行了展望。



Abstract  

Synthetic biology is an interdisciplinary field that applies engineering principles to design and construct novel biological systems or organisms. Initially focused on microbial systems, its applications have expanded to include plants. Plant synthetic biology offers promising solutions to pressing global challenges in agriculture and human health. As a staple crop for much of the world’s population and a model species in plant science, rice has emerged as a pivotal platform in this domain. Significant progress has been achieved in genome engineering through multiplex genome editing, synthetic hybrid rice systems, induction of apomixis, reconstruction of photosynthesis and nitrogen-fixation pathways, and biosynthesis of micronutrients, pharmaceuticals, and therapeutic proteins or peptides. This review summarizes recent advances in rice synthetic biology, outlines current developments, and discusses future research directions.

Keywords:  rice synthetic biology       multiplex genome editing       synthetic hybrid rice       engineered pathways       endosperm bioreactor  
Online: 22 December 2025  
Fund: 

We acknowledge the financial support from the Zhejiang Provincial Natural Science Foundation of China (LD24C130002 and LQN25C020001) and the National Natural Science Foundation of China (32400208).

About author:  Chenchen Zhang, E-mail: zcshiyu@163.com; #Sheng Teng, E-mail: steng@cjlu.edu.cn

Cite this article: 

Chenchen Zhang, Yan Wang, Lu Chen, Xixi Wang, Sheng Teng. 2025. Advances in rice synthetic biology: toward a better staple crop and beyond. Journal of Integrative Agriculture, Doi:10.1016/j.jia.2025.12.036

Abramson J, Adler J, Dunger J, Evans R, Green T, Pritzel A, Ronneberger O, Willmore L, Ballard A J, Bambrick J, Bodenstein S W, Evans D A, Hung C C, O’neill M, Reiman D, Tunyasuvunakool K, Wu Z, Žemgulytė A, Arvaniti E, Beattie C, et al. 2024. Accurate structure prediction of biomolecular interactions with AlphaFold 3. Nature, 630, 493-500.

Alappat B, Alappat J. 2020. Anthocyanin pigments: Beyond aesthetics. Molecules, 25, 5500.

Albertsen M C, Fox T W, Hershey H P. 2006. Nucleotide Sequences Mediating Plant Male Fertility and Method of Using Same. U.S. Patent, Patent No. WO2007002267, Patent and Trademark Office, Washington, D.C., USA.

De Almeida B P, Richard G, Dalla-Torre H, Blum C, Hexemer L, Pandey P, Laurent S, Rajesh C, Lopez M, Laterre A, Lang M, Şahin U, Beguir K, Pierrot T. 2025. A multimodal conversational agent for DNA, RNA and protein tasks. Nature Machine Intelligence, 7, 928-941.

Aluko R E. 2015. Antihypertensive peptides from food proteins. Annual Review of Food Science and Technology, 6, 235-262.

An N, Ou J, Jiang D, Zhang L, Liu J, Fu K, Dai Y, Yang D. 2013. Expression of a functional recombinant human basic fibroblast growth factor from transgenic rice seeds. International Journal of Molecular Sciences, 14, 3556-3567.

An X, Dong Z, Tian Y, Xie K, Wu S, Zhu T, Zhang D, Zhou Y, Niu C, Ma B, Hou Q, Bao J, Zhang S, Li Z, Wang Y, Yan T, Sun X, Zhang Y, Li J, Wan X. 2019. ZmMs30 encoding a novel GDSL lipase is essential for male fertility and valuable for hybrid breeding in maize. Molecular Plant, 12, 343-359.

An X, Ma B, Duan M, Dong Z, Liu R, Yuan D, Hou Q, Wu S, Zhang D, Liu D, Yu D, Zhang Y, Xie K, Zhu T, Li Z, Zhang S, Tian Y, Liu C, Li J, Yuan L, et al. 2020. Molecular regulation of ZmMs7 required for maize male fertility and development of a dominant male-sterility system in multiple species. Proceedings of the National Academy of Sciences of the United States of America, 117, 23499-23509.

Anzalone A V, Koblan L W, Liu D R. 2020. Genome editing with CRISPR–Cas nucleases, base editors, transposases and prime editors. Nature Biotechnology, 38, 824-844.

Baek S H, Shin W C, Ryu H S, Lee D W, Moon E, Seo C S, Hwang E, Lee H S, Ahn M H, Jeon Y, Kang H J, Lee S W, Kim S Y, D’souza R, Kim H J, Hong S T, Jeon J S. 2013. Creation of resveratrol-enriched rice for the treatment of metabolic syndrome and related diseases. PLoS ONE, 8, e57930.

Bai C, Berman J, Farre G, Capell T, Sandmann G, Christou P, Zhu C. 2017. Reconstruction of the astaxanthin biosynthesis pathway in rice endosperm reveals a metabolic bottleneck at the level of endogenous β-carotene hydroxylase activity. Transgenic Research, 26, 13-23.

Bao J, Zhou X, Xu F, He Q, Park Y J. 2017. Genome-wide association study of the resistant starch content in rice grains. Starch-Stärke, 69, 1600343.

Baron F, Nagler A. 2017. Novel strategies for improving hematopoietic reconstruction after allogeneic hematopoietic stem cell transplantation or intensive chemotherapy. Expert Opinion on Biological Therapy, 17, 163-174.

Bekker L G, Beyrer C, Mgodi N, Lewin S R, Delany-Moretlwe S, Taiwo B, Masters M C, Lazarus J V. 2023. HIV infection. Nature Reviews Disease Primers, 9, 42.

Blancquaert D, Van Daele J, Strobbe S, Kiekens F, Storozhenko S, De Steur H, Gellynck X, Lambert W, Stove C, Van Der Straeten D. 2015. Improving folate (vitamin B9) stability in biofortified rice through metabolic engineering. Nature Biotechnology, 33, 1076-1078.

Bradbury A, Clapp O, Biacsi A S, Kuo P, Gaju O, Hayta S, Zhu J K, Lambing C. 2025. Integrating genome editing with omics, artificial intelligence and advanced farming technologies to increase crop productivity. Plant Communications, 101386.

Calzada J P V, Crane C F, Stelly D M. 1996. Apomixis-the asexual revolution. Science, 274, 1322-1323.

Chai N, Xu J, Zhang R, Li G, Wen J, Su L, Xue Y, Li T, Liu J, Zeng D, Tan J, Huang J, Chen L, Liu Y G, Zhu Q. 2025. Synthetic metabolic engineering of functional crops: Boosting nutrition and human health. The Crop Journal, doi: 10.1016/j.xplc.2019.100017.

Chang Z, Chen Z, Wang N, Xie G, Lu J, Yan W, Zhou J, Tang X, Deng X W. 2016. Construction of a male sterility system for hybrid rice breeding and seed production using a nuclear male sterility gene. Proceedings of the National Academy of Sciences of the United States of America, 113, 14145-14150.

Chen G, Li Y, Jin K, Gao J, Wu S, Cui X, Mao C, Yin X, Lu T, Zhang Z. 2025. Synthetic photorespiratory bypass improves rice productivity by enhancing photosynthesis and nitrogen uptake. The Plant Cell, 37, koaf015.

Chen J H, Chen S T, He N Y, Wang Q L, Zhao Y, Gao W, Guo F Q. 2020. Nuclear-encoded synthesis of the D1 subunit of photosystem II increases photosynthetic efficiency and crop yield. Nature Plants, 6, 570-580.

Chen K, Sun K, Ye C, Liu Y, Hu Y, Jia B, Jiang S, Zou Y, Cao R, Guo J, Chen D, Zhou X, Liu J, Qian Q, Peng X, Shen M, Zhou D, Cheng S, Mi J, Zhu Q, et al. 2025. Metabolic engineering of theanine biosynthesis in rice endosperm to develop biofortified theaRice for benefiting human health. Plant Communications, 101411.

Chen R, Deng Y, Ding Y, Guo J, Qiu J, Wang B, Wang C, Xie Y, Zhang Z, Chen J, Chen L, Chu C, He G, He Z, Huang X, Xing Y, Yang S, Xie D, Liu Y, Li J. 2022. Rice functional genomics: Decades’ efforts and roads ahead. Science China Life Sciences, 65, 33-92.

Chen X, Li G, He H, Xie W, Cui L, Zhang Z, Peng X, Zhu G. 2025. A synthetic glycolate metabolism bypass in rice chloroplasts increases photosynthesis and yield. The Crop Journal, doi.org/10.1016/j.cj.2025.03.001.

Chen Z, Du H, Tao Y, Xu Y, Wang F, Li B, Zhu Q H, Niu H, Yang J. 2022. Efficient breeding of low glutelin content rice germplasm by simultaneous editing multiple glutelin genes via CRISPR/Cas9. Plant Science, 324, 111449.

Chen Z, Wu J, Deng X W, Tang X. 2023. Establishment and advances of third-generation hybrid rice technology: A review. Rice, 16, 56.

Cheng T, Sun M X. 2025. A highly efficient and environmentally friendly strategy to obtain transgene-free genome-edited progeny in flowering plants. Molecular Plant, 18, 911-914.

Dan J, Xia Y, Wang Y, Zhan Y, Tian J, Tang N, Deng H, Cao M. 2024. One-line hybrid rice with high-efficiency synthetic apomixis and near-normal fertility. Plant Cell Reports, 43, 79.

Dash R, Bhattacharjya S. 2021. Thanatin: An emerging host defense antimicrobial peptide with multiple modes of action. International Journal of Molecular Sciences, 22, 1522.

Deng X, Wang H, Tang X, Zhou J, Chen H, He G, Chen L, Xu Z. 2013. Hybrid rice breeding welcomes a new era of molecular crop design. Scientia Sinica Vitae, 43, 864-868. (in Chinese)

D'Erfurth I, Jolivet S, Froger N, Catrice O, Novatchkova M, Mercier R. 2009. Turning meiosis into mitosis. Plos Biology, 7, e1000124.

Dong M, Masuyer G, Stenmark P. 2019. Botulinum and tetanus neurotoxins. Annual Review of Biochemistry, 88, 811-837.

Dong X X, Jin H L, Wang H B. 2016. Research progress on adaptive mechanisms of photosystem to highlight in plants. Plant Physiology Journal, 52, 1725-32. (in Chinese)

Du S, Wang Z, Chen Y, Tan Y, Li X, Zhu W, He G, Lei K, Guo L, Zhang Y. 2022. Coleoptile purple line regulated by A-P gene system is a valuable marker trait for seed purity identification in hybrid rice. Rice Science, 29, 451-461.

Fischer T K, Viboud C, Parashar U, Malek M, Steiner C, Glass R, Simonsen L. 2007. Hospitalizations and deaths from diarrhea and rotavirus among children <5 years of age in the United States, 1993–2003. The Journal of Infectious Diseases, 195, 1117-1125.

Flotte T R, Brantly M L, Spencer L T, Byrne B J, Spencer C T, Baker D J, Humphries M. 2004. Phase I trial of intramuscular injection of a recombinant adeno-associated virus alpha 1-antitrypsin (rAAV2-CB-hAAT) gene vector to AAT-deficient adults. Human Gene Therapy, 15, 93-128.

Fujiwara Y, Aiki Y, Yang L, Takaiwa F, Kosaka A, Tsuji N M, Shiraki K, Sekikawa K. 2010. Extraction and purification of human interleukin-10 from transgenic rice seeds. Protein Expression and Purification, 72, 125-130.

Gannon B M, Huey S L, Mehta N H, Shrestha N, Lopez-Perez L, Martinez R X, Rogers L M, Garcia-Casal M N, Mehta S. 2025. Selected laboratory‐based biomarkers for assessing vitamin A deficiency in at risk individuals. Cochrane Database of Systematic Reviews, 5, CD013742.

Ge J, Lang X, Ji J, Qu C, Qiao H, Zhong J, Luo D, Hu J, Chen H, Wang S, Wang T, Li S, Li W, Zheng P, Xu J, Du H. 2024. Integration of biological and information technologies to enhance plant autoluminescence. The Plant Cell, 36, 4703-4715.

Guo K, Yang J, Yu N, Luo L, Wang E. 2023. Biological nitrogen fixation in cereal crops: Progress, strategies, and perspectives. Plant Commun, 4, 100499.

Ha S H, Kim J K, Jeong Y S, You M K, Lim S H, Kim J K. 2019. Stepwise pathway engineering to the biosynthesis of zeaxanthin, astaxanthin and capsanthin in rice endosperm. Metabolic Engineering, 52, 178-189.

Ha S H, Liang Y S, Jung H, Ahn M J, Suh S C, Kweon S J, Kim D H, Kim Y M, Kim J K. 2010. Application of two bicistronic systems involving 2A and IRES sequences to the biosynthesis of carotenoids in rice endosperm. Plant Biotechnology Journal, 8, 928-938.

Han J Y, Baek S H, Jo H J, Yun D W, Choi Y E. 2019. Genetically modified rice produces ginsenoside aglycone (protopanaxadiol). Planta, 250, 1103-1110.

Hao Y, Xu B, Zong W, Li S, Du D, Chen M, Xiao D, Song Y, Guo X, Li W, Wu Z, Zhang K, Liao N, Hu D, Liu Y G, Guo J. 2025. Directional improvement of agronomic traits in salt-tolerant rice by multiplex-genome-editing. Journal of Integrative Plant Biologydoi: 10.1111/jipb.13926.

Hathwaik L T, Horstman J, Thomson J G, Thilmony R. 2021. Efficient gene stacking in rice using the GAANTRY system. Rice, 14, 17.

He W, Wang L, Lin Q, Yu F. 2021. Rice seed storage proteins: Biosynthetic pathways and the effects of environmental factors. Journal of Integrative Plant Biology, 63, 1999-2019.

He Y, Fang P, Shan Y, Pan Y, Wei Y, Chen Y, Chen Y, Liu Y, Zeng Z, Zhou Z, Zhu F, Holmes E C, Ye J, Li J, Shu Y, Shi M, Li Z. 2025. Generalized biological foundation model with unified nucleic acid and protein language. Nature Machine Intelligence, 7, 942-953.

He Y, Ning T, Xie T, Qiu Q, Zhang L, Sun Y, Jiang D, Fu K, Yin F, Zhang W, Shen L, Wang H, Li J, Lin Q, Sun Y, Li H, Zhu Y, Yang D. 2011. Large-scale production of functional human serum albumin from transgenic rice seeds. Proceedings of the National Academy of Sciences of the United States of America, 108, 19078-19083.

Hu F, Liu C, Jin X, Sun T, Hong L, Rao Y, Qian Q, Wang K. 2025. OsPLDα2-dependent synthetic apomixis enables normal seed setting in hybrid rice via genome editing. Science Bulletin, doi: 10.1016/j.scib.2025.05.022.

Hu S, Zhou L, Wang J, Mawia A M, Hui S, Xu B, Jiao G, Sheng Z, Shao G, Wei X, Wang L, Xie L, Zhao F, Tang S, Hu P. 2024. Production of grains with ultra-low heavy metal accumulation by pyramiding novel Alleles of OsNramp5 and OsLsi2 in two-line hybrid rice. Plant Biotechnology Journal, 22, 2921-2931.

Huang J, Nandi S, Wu L, Yalda D, Bartley G, Rodriguez R, Lonnerdal B, Huang N. 2002. Expression of natural antimicrobial human lysozyme in rice grains. Molecular Breeding, 10, 83-94.

Huang Y, Dong H, Shang M, Wang K. 2021. CRISPR/Cas systems: The link between functional genes and genetic improvement. The Crop Journal, 9, 678-687.

Huang Y, Meng X, Rao Y, Xie Y, Sun T, Chen W, Wei X, Xiong J, Yu H, Li J, Wang K. 2025. OsWUS-driven synthetic apomixis in hybrid rice. Plant Communications, 6, 101136.

Huang Z, Lin J, Cheng Z, Xu M, Guo M, Huang X, Yang Z, Zheng J. 2015a. Production of oleanane-type sapogenin in transgenic rice via expression of β-amyrin synthase gene from Panax japonicus C. A. Mey. BMC Biotechnology, 15, 45.

Huang Z, Lin J, Cheng Z, Xu M, Huang X, Yang Z, Zheng J. 2015b. Production of dammarane-type sapogenins in rice by expressing the dammarenediol-II synthase gene from Panax ginseng C.A. Mey. Plant Science, 239, 106-114.

Hong W J, Kim Y J, Chandran A K N, Jung K H. 2019. Infrastructures of systems biology that facilitate functional genomic study in rice. Rice, 12, 15.

Isemura M. 2019. Catechin in human health and disease. Molecules, 24, 528.

Iyer R, Tomar S K. 2009. Folate: A functional food constituent. Journal of Food Science, 74, R114-R122.

Jeong Y S, Ku H K, Kim J K, You M K, Lim S H, Kim J K, Ha S H. 2017. Effect of codon optimization on the enhancement of the β-carotene contents in rice endosperm. Plant Biotechnology Reports, 11, 171-179.

Jollès P. 1964. Recent developments in the study of lysozymes. Angewandte Chemie International Edition in English, 3, 28-36.

Khanday I, Skinner D, Yang B, Mercier R, Sundaresan V. 2019. A male-expressed rice embryogenic trigger redirected for asexual propagation through seeds. Nature, 565, 91-95.

Koltunow A M, Grossniklaus U. 2003. Apomixis: A developmental perspective. Annual Review of Plant Biology, 54, 547-574.

Kumrungsee T, Peipei zhang, Yanaka N, Suda T, Kato N. 2022. Emerging cardioprotective mechanisms of vitamin B6: A narrative review. European Journal of Nutrition, 61, 605-613.

Lai C, Hu S, Jiao G, Wang L, Shao G, Zhao F, Xie L, Wei X, Lyu Y, Sheng Z, Tang S, Hu P. 2025. Enhancing folate content in japonica rice through co-expression of OsADCS and OsGTPCHI indica alleles. Rice Science, 32, 353-366.

Lee C. 2019. Griffithsin, a highly potent broad-spectrum antiviral lectin from red algae: From discovery to clinical application. Marine Drugs, 17, 567.

Lee D S, Chen L J, Li C Y, Liu Y, Tan X L, Lu B R, Li J, Gan S X, Kang S G, Suh H S, Zhu Y. 2013. The bsister MADS gene FST determines ovule patterning and development of the zygotic embryo and endosperm. PLoS ONE, 8, e58748.

Lei D, Tang W, Xie Z Liu H, Chen LY. 2013. How to tackle insecurity problems in seed production of two-line hybrid rice. Acta Agronomica Sinica, 39,1569-1575. (in Chinese)

Lee D Y, Noren Hooten N, O'connell J F, Lee B Y, Kim Y. 2025. The role of ginseng and its bioactive compounds in aging: Cells and animal studies. Annual Review of Food Science and Technology, 16, 333-354.

Leister D. 2023. Enhancing the light reactions of photosynthesis: Strategies, controversies, and perspectives. Molecular Plant, 16, 4-22.

Li H, Pinot F, Sauveplane V, Werck-Reichhart D, Diehl P, Schreiber L, Franke R, Zhang P, Chen L, Gao Y, Liang W, Zhang D. 2010. Cytochrome P450 family member CYP704B2 catalyzes the ω-hydroxylation of fatty acids and is required for anther cutin biosynthesis and pollen exine formation in rice. The Plant Cell, 22, 173-190.

Li M Y, Liu H Y, Wu D T, Kenaan A, Geng F, Li H B, Gunaratne A, Li H, Gan R Y. 2022. L-theanine: A unique functional amino acid in tea (Camellia sinensis L.) with multiple health benefits and food applications. Frontiers in Nutrition, 9, 853846.

Li S, Li W, Huang B, Cao X, Zhou X, Ye S, Li C, Gao F, Zou T, Xie K, Ren Y, Ai P, Tang Y, Li X, Deng Q, Wang S, Zheng A, Zhu J, Liu H, Wang L, et al. 2013. Natural variation in PTB1 regulates rice seed setting rate by controlling pollen tube growth. Nature Communications, 4, 2793.

Li W, Guo X, Wu W, Yu W, Li S, Luo D, Wang T, Zhu Q, Chen L, Lee D. 2022. Construction of a novel female sterility system for hybrid rice. Frontiers in Plant Science, 12, 815401.

Li Y L, Tang H Y, Huang Q C, Li X Q. 2024. Research and application of the third-generation hybrid rice. Molecular Plant Breeding, 22, 5124-5131 (in Chinese).

Liao C, Yan W, Chen Z, Xie G, Deng X W, Tang X. 2021. Innovation and development of the third-generation hybrid rice technology. The Crop Journal, 9, 693-701.

Liu C, He Z, Zhang Y, Hu F, Li M, Liu Q, Huang Y, Wang J, Zhang W, Wang C, Wang K. 2023. Synthetic apomixis enables stable transgenerational transmission of heterotic phenotypes in hybrid rice. Plant Communications, 4, 100470.

Liu J, Zhang R, Chai N, Su L, Zheng Z, Liu T, Guo Z, Ma Y, Xie Y, Xie X, Lin Q, Chen L, Liu Y G, Zhu Q. 2025. Programmable genome engineering and gene modifications for plant biodesign. Plant Communications, 6, 101427.

Liu P, He L, Mei L, Zhai W, Chen X, Ma B. 2022. Rapid and directional improvement of elite rice variety via combination of genomics and multiplex genome editing. Journal of Agricultural and Food Chemistry, 70, 6156-6167.

Liu X, Li Z, Ying J, Shu Y, Liu W, Li G, Chen L, Luo J, Wang S, Wang Y, Tong X, Huang J, Du H, Zhang J. 2024. Multi-gene engineering boosts oil content in rice grains. Plant Commun, 5, 100736.

Lu H P, Liu S M, Xu S L, Chen W Y, Zhou X, Tan Y Y, Huang J Z, Shu Q Y. 2017. CRISPR-S: An active interference element for a rapid and inexpensive selection of genome-edited, transgene-free rice plants. Plant Biotechnology Journal, 15, 1371-1373.

Majumder S, Datta K, Datta S K. 2019. Rice biofortification: High iron, zinc, and vitamin-A to fight against “Hidden Hunger”. Agronomy, 9, 803.

Makarchikov A F, Wins P, Bettendorff L. 2025. Biochemical and medical aspects of vitamin B1 research. Neurochemistry International, 185, 105962.

Mangel N, Fudge J B, Li K T, Wu T Y, Tohge T, Fernie A R, Szurek B, Fitzpatrick T B, Gruissem W, Vanderschuren H. 2019. Enhancement of vitamin B6 levels in rice expressing Arabidopsis vitamin B6 biosynthesis de novo genes. The Plant Journal, 99, 1047-1065.

Mares J. 2016. Lutein and zeaxanthin isomers in eye health and disease. Annual Review of Nutrition, 36, 571-602.

Matsumoto S, Tanaka R, Okada K, Arita K, Hyakusoku H, Miyamoto M, Tabata Y, Mizuno H. 2013. The effect of control-released basic fibroblast growth factor in wound healing: Histological analyses and clinical application. Plastic and Reconstructive SurgeryGlobal Open, 1, e44.

Matsumoto Y, Suzuki S, Nozoye T, Yamakawa T, Takashima Y, Arakawa T, Tsuji N, Takaiwa F, Hayashi Y. 2009. Oral immunogenicity and protective efficacy in mice of transgenic rice plants producing a vaccine candidate antigen (As16) of Ascaris suum fused with cholera toxin B subunit. Transgenic Research, 18, 185-192.

Miao C, Xiao L, Hua K, Zou C, Zhao Y, Bressan R A, Zhu J K. 2018. Mutations in a subfamily of abscisic acid receptor genes promote rice growth and productivity. Proceedings of the National Academy of Sciences of the United States of America, 115, 6058-6063.

Mieulet D, Jolivet S, Rivard M, Cromer L, Vernet A, Mayonove P, Pereira L, Droc G, Courtois B, Guiderdoni E, Mercier R. 2016. Turning rice meiosis into mitosis. Cell Research, 26, 1242-1254.

Mo B, Chen X, Yang J, Chen L, Guo W, Wu S, Peng X, Zhang Z. 2025. Engineering of photorespiration-dependent glycine betaine biosynthesis improves photosynthetic carbon fixation and panicle architecture in rice. Journal of Integrative Plant Biology, 67, 979-992.

Monmai C, Baek S H. 2024. Anti-inflammatory effects of the combined treatment of resveratrol- and protopanaxadiol-enriched rice seed extract on lipopolysaccharide-stimulated RAW264.7 cells. Molecules, 29, 4343.

Monmai C, Kim J S, Baek S H. 2022. Transgenic rice seed extracts exert immunomodulatory effects by modulating immune-related biomarkers in RAW264.7 macrophage cells. Nutrients, 14, 4143.

Monmai C, Kim J S, Baek S H. 2023. Effect of ginseng sapogenin protopanaxadiol-enriched rice (DJ-PPD) on immunomodulation. Plants, 12, 767.

Moore K W, De Waal Malefyt R, Coffman R L, Apos, Garra A. 2001. Interleukin-10 and the interleukin-10 receptor. Annual Review of Immunology, 19, 683-765.

Nandi S, Suzuki Y A, Huang J, Yalda D, Pham P, Wu L, Bartley G, Huang N, Lönnerdal B. 2002. Expression of human lactoferrin in transgenic rice grains for the application in infant formula. Plant Science, 163, 713-722.

Nandi S, Yalda D, Lu S, Nikolov Z, Misaki R, Fujiyama K, Huang N. 2005. Process development and economic evaluation of recombinant human lactoferrin expressed in rice grain. Transgenic Res, 14, 237-249.

Ning T, Xie T, Qiu Q, Yang W, Zhou S, Zhou L, Zheng C, Zhu Y, Yang D. 2008. Oral administration of recombinant human granulocyte-macrophage colony stimulating factor expressed in rice endosperm can increase leukocytes in mice. Biotechnology Letters, 30, 1679-1686.

Niu J, Gao Y, Wang G, Qin Z, Wu C, Yu Z, Wang L, Hu Z, Li X, Zhang Z, Chen Y, Yao L, Yang J, Li G M, Yang Y, Lu X, Gu Y, Wu X, Mao X, Zhou Z, et al. 2025. Rice-derived recombinant human serum albumin as an alternative to human plasma for patients with decompensated liver cirrhosis: A randomised, double-blind, positive-controlled and non-inferiority trial. Gut, 74, 1476-1485.

Nochi T, Takagi H, Yuki Y, Yang L, Masumura T, Mejima M, Nakanishi U, Matsumura A, Uozumi A, Hiroi T, Morita S, Tanaka K, Takaiwa F, Kiyono H. 2007. Rice-based mucosal vaccine as a global strategy for cold-chain- and needle-free vaccination. Proceedings of the National Academy of Sciences of the United States of America, 104, 10986-10991.

Nojima J, Ishii-Katsuno R, Futai E, Sasagawa N, Watanabe Y, Yoshida T, Ishiura S. 2011a. Production of anti-amyloid β antibodies in mice fed rice expressing amyloid β. Bioscience, Biotechnology, and Biochemistry, 75, 396-400.

Nojima J, Maeda A, Aoki S, Suo S, Yanagihara D, Watanabe Y, Yoshida T, Ishiura S. 2011b. Effect of rice-expressed amyloid β in the Tg2576 Alzheimer’s disease transgenic mouse model. Vaccine, 29, 6252-6258.

Ogo Y, Ozawa K, Ishimaru T, Murayama T, Takaiwa F. 2013. Transgenic rice seed synthesizing diverse flavonoids at high levels: A new platform for flavonoid production with associated health benefits. Plant Biotechnology Journal, 11, 734-746.

Oliva N, Chadha-Mohanty P, Poletti S, Abrigo E, Atienza G, Torrizo L, Garcia R, Dueñas C, Poncio M A, Balindong J, Manzanilla M, Montecillo F, Zaidem M, Barry G, Hervé P, Shou H, Slamet-Loedin I H. 2014. Large-scale production and evaluation of marker-free indica rice IR64 expressing phytoferritin genes. Molecular Breeding, 33, 23-37.

Oliva R, Ji C, Atienza-Grande G, Huguet-Tapia J C, Perez-Quintero A, Li T, Eom J S, Li C, Nguyen H, Liu B, Auguy F, Sciallano C, Luu V T, Dossa G S, Cunnac S, Schmidt S M, Slamet-Loedin I H, Vera Cruz C, Szurek B, Frommer W B, et al. 2019. Broad-spectrum resistance to bacterial blight in rice using genome editing. Nature Biotechnology, 37, 1344-1350.

Oszvald M, Kang T J, Tomoskozi S, Jenes B, Kim T G, Cha Y S, Tamas L, Yang M S. 2008. Expression of cholera toxin B subunit in transgenic rice endosperm. Molecular Biotechnology, 40, 261-268.

Ozias-Akins P, Van Dijk P J. 2007. Mendelian genetics of apomixis in plants. Annual Review of Genetics, 41, 509-537.

Paine J A, Shipton C A, Chaggar S, Howells R M, Kennedy M J, Vernon G, Wright S Y, Hinchliffe E, Adams J L, Silverstone A L, Drake R. 2005. Improving the nutritional value of Golden Rice through increased pro-vitamin A content. Nature Biotechnology, 23, 482-487.

Papikian A, Liu W, Gallego-Bartolomé J, Jacobsen S E. 2019. Site-specific manipulation of Arabidopsis loci using CRISPR-Cas9 SunTag systems. Nature Communications, 10, 729.

Patel-Tupper D, Kelikian A, Leipertz A, Maryn N, Tjahjadi M, Karavolias N G, Cho M J, Niyogi K K. 2024. Multiplexed CRISPR-Cas9 mutagenesis of rice PSBS1 noncoding sequences for transgene-free overexpression. Science Advances, 10, eadm7452.

Peng F, Li C, Lu C, Li Y, Xu P, Liang G. 2022. IRONMAN peptide interacts with OsHRZ1 and OsHRZ2 to maintain Fe homeostasis in rice. Journal of Experimental Botany, 73, 6463-6474.

Perez-Prat E, Van Lookeren Campagne M M. 2002. Hybrid seed production and the challenge of propagating male-sterile plants. Trends in Plant Science, 7, 199-203.

Prodhan Z H, Shu Q Y. 2020. Rice aroma: A natural gift comes with price and the way forward. Rice Science, 27, 86-100.

Qian D, Qiu B, Zhou N, Takaiwa F, Yong W, Qu L Q. 2020. Hypotensive activity of transgenic rice seed accumulating multiple antihypertensive peptides. Journal of Agricultural and Food Chemistry, 68, 7162-7168.

Rahimi P, Saeed A, Alireza M N S, Ostadrahimi A. 2019. Betalains, the nature-inspired pigments, in health and diseases. Critical Reviews in Food Science and Nutrition, 59, 2949-2978.

Ren H, Shankle K, Cho M J, Tjahjadi M, Khanday I, Sundaresan V. 2024. Synergistic induction of fertilization-independent embryogenesis in rice egg cells by paternal-genome-expressed transcription factors. Nature Plants, 10, 1892-1899.

Rezaei E E, Webber H, Asseng S, Boote K, Durand J L, Ewert F, Martre P, Maccarthy D S. 2023. Climate change impacts on crop yields. Nature Reviews Earth & Environment, 4, 831-846.

Saeed P A, Ghorbani A, Rostami M, Ashrafi-Dehkordi E. 2025. Integrating multi-omic data to identify key genes and pathways involved in rice bacterial leaf streak disease. Scientific Reports, 15, 21025.

Sang S, Sun X, Ma T, Zhang Y, Yao G, Wang X, Tan X, Feng L, Li J, Ji S, Cheng H. 2025. Efficient promoter editing of the SBEIIb gene enables fine-tuning of the resistant starch content in rice. International Journal of Biological Macromolecules, 290, 138904.

Sardana R, Dudani A K, Tackaberry E, Alli Z, Porter S, Rowlandson K, Ganz P, Altosaar I. 2007. Biologically active human GM-CSF produced in the seeds of transgenic rice plants. Transgenic Research, 16, 713-721.

Sasaki T, International Rice Genome Sequencing P. 2005. The map-based sequence of the rice genome. Nature, 436, 793-800.

Schenk D, Barbour R, Dunn W, Gordon G, Grajeda H, Guido T, Hu K, Huang J, Johnson-Wood K, Khan K, Kholodenko D, Lee M, Liao Z, Lieberburg I, Motter R, Mutter L, Soriano F, Shopp G, Vasquez N, Vandevert C, et al. 1999. Immunization with amyloid-β attenuates Alzheimer-disease-like pathology in the PDAPP mouse. Nature, 400, 173-177.

Senguttuvel P, Padmavathi G, Jasmine C, Sanjeeva Rao D, Neeraja C, Jaldhani V, Beulah P, Gobinath R, Aravind Kumar J, Sai Prasad S, Subba Rao L, Hariprasad A, Sruthi K, Shivani D, Sundaram R, Govindaraj M. 2023. Rice biofortification: Breeding and genomic approaches for genetic enhancement of grain zinc and iron contents. Frontiers in Plant Science, 14, 1138408.

Shang Y, Shi H, Liu M, Lan P, Li D, Liu X, Wang M, Zhang Z, Chen S. 2025. Using synthetic biology to express nitrogenase biosynthesis pathway in rice and to overcome barriers of nitrogenase instability in plant cytosol. Trends in Biotechnology, 43, 946-968.

Shen B R, Wang L M, Lin X L, Yao Z, Xu H W, Zhu C H, Teng H Y, Cui L L, Liu E E, Zhang J J, He Z H, Peng X X. 2019. Engineering a new chloroplastic photorespiratory bypass to increase photosynthetic efficiency and productivity in rice. Molecular Plant, 12, 199-214.

Shen N, Wang T, Gan Q, Liu S, Wang L, Jin B. 2022. Plant flavonoids: Classification, distribution, biosynthesis, and antioxidant activity. Food Chemistry, 383, 132531.

Shin Y M, Park H J, Yim S D, Baek N I, Lee C H, An G, Woo Y M. 2006. Transgenic rice lines expressing maize C1 and R-S regulatory genes produce various flavonoids in the endosperm. Plant Biotechnology Journal, 4, 303-315.

Singh R, Kaur N, Praba U P, Kaur G, Tanin M J, Kumar P, Neelam K, Sandhu J S, Vikal Y. 2022. A prospective review on selectable marker-free genome engineered rice: Past, present and future scientific realm. Frontiers in Genetics, 13, 882836.

Sixma T K, Pronk S E, Kalk K H, Wartna E S, Van Zanten B A M, Witholt B, Hoi W G J. 1991. Crystal structure of a cholera toxin-related heat-labile enterotoxin from E. coli. Nature, 351, 371-377.

Song L, Wen Y, Zhou Y, Zhang H, Tian Y, Wang J, Cui Y, Tan R, Xiong D, Meng C, Zhou Y, Li Q, Pan Z, Liu Q, Jiao X. 2025. Rice-derived SARS-CoV-2 glycoprotein S1 subunit vaccine elicits humoral and cellular immune responses. Plant Biotechnology Journal, 23, 2570-2582

Song M, Li F, Chen Z, Hou H, Wang Y, Liu H, Liu D, Li J, Peng T, Zhao Y, Du Y, Sun H, Du C, Zhang J, Zhao Q, Miao C. 2024a. Engineering high-frequency apomixis with normal seed production in hybrid rice. iScience, 27, 111479.

Song M, Wang W, Ji C, Li S, Liu W, Hu X, Feng A, Ruan S, Du S, Wang H, Dai K, Guo L, Qian Q, Si H, Hu X. 2024b. Simultaneous production of high-frequency synthetic apomixis with high fertility and improved agronomic traits in hybrid rice. Molecular Plant, 17, 4-7.

Song S, Wang T, Li Y, Hu J, Kan R, Qiu M, Deng Y, Liu P, Zhang L, Dong H, Li C, Yu D, Li X, Yuan D, Yuan L, Li L. 2021. A novel strategy for creating a new system of third-generation hybrid rice technology using a cytoplasmic sterility gene and a genic male-sterile gene. Plant Biotechnology Journal, 19, 251-260.

Song X, Qiao Y, Ma J, Zhang X, Liu J, Xin W, Xing S, Wang Y. 2023. Co-expression of four penaeidins in transgenic rice seeds: An alternative strategy for substitute antibiotic agricultural products. Transgenic Research, 32, 463-473.

Spagnoli A, Longobardi L, O’rear L. 2005. Cartilage disorders: Potential therapeutic use of mesenchymal stem cells. Endocrine Development, 9, 17-30.

Stompor M. 2020. A review on sources and pharmacological aspects of sakuranetin. Nutrients, 12, 513.

Storozhenko S, De Brouwer V, Volckaert M, Navarrete O, Blancquaert D, Zhang G F, Lambert W, Van Der Straeten D. 2007. Folate fortification of rice by metabolic engineering. Nature Biotechnology, 25, 1277-1279.

Strobbe S, Verstraete J, Stove C, Van Der Straeten D. 2021. Metabolic engineering of rice endosperm towards higher vitamin B1 accumulation. Plant Biotechnology Journal, 19, 1253-1267.

Susaki D, Suzuki T, Maruyama D, Ueda M, Higashiyama T, Kurihara D. 2021. Dynamics of the cell fate specifications during female gametophyte development in Arabidopsis. Plos Biology, 19, e3001123.

Tan J, Sun K, Qian Q, Chai N, Peng X, Shen M, Zhou D, Liu Y G, Zhu Q, Liu Q. 2025a Development of PAM-flexible multiplex genome editors for enhancing cold-tolerance in indica rice. The Crop Journal, 13, 991-995.

Tan J, Wang Y, Lin Z, Chai N, Xue Y, Chen L, Liu Y G, Zhu Q. 2025b eRUBY rice: Co-expression of a feedback-insensitive TyrA arogenate dehydrogenase with RUBY enhances endosperm betalain levels. Plant Physiology, 199, kiaf416.

Tan S, Han R, Li P, Yang G, Li S, Zhang P, Wang W B, Zhao W Z, Yin L P. 2015. Over-expression of the MxIRT1 gene increases iron and zinc content in rice seeds. Transgenic Research, 24, 109-122.

Tan W, Liang S, Gu X, Yi C, Liang G, Zhang C, Zhou Y. 2024. Application and prospects of genome-editing technology for improvement of grain yield and quality in rice. Chinese Science Bulletin, 70, 2468-2482.

Tanenbaum M E, Gilbert L A, Qi L S, Weissman J S, Vale R D. 2014. A protein-tagging system for signal amplification in gene expression and fluorescence imaging. Cell, 159, 635-646.

Tao H, Xiao N, Wang R, He F, Cai Y, Jiang S, Wang M, Wang D, Chen H, You X, Li A, Wang G L, Ning Y. 2025. Development of elite rice with broad-spectrum resistance through pyramiding of key resistance gene and simultaneously editing multiple susceptibility genes. Journal of Integrative Plant Biology, doi: 10.1111/jipb.13901.

Tao Y, Chen Z, Xu Y, Wang F, Jiang Y, Fan F, Li W, Zhu J, Li X, Wang J, Zhu Q H, Xu G, Yang J. 2025. CRISPR-mediated targeted mutagenesis for improving nitrogen use efficiency in japonica rice. Plant Communications, 6, 101189.

Teng B, Chen Z, Ying Z, Shiyun D, Min X, Fengshun S, Jinlong N, Zhixiang L, And Ni D. 2018. Effects of different Wx alleles on amylopectin molecular structure and enzymatic hydrolysis properties of rice starch. International Journal of Food Properties, 21, 2772-2784.

Thakur K, Kumar T S, Kumar S A, Surajit M, And Arora S. 2017. Riboflavin and health: A review of recent human research. Critical Reviews in Food Science and Nutrition, 57, 3650-3660.

Tian H, Li Y, Lu Y, Zhang Q, Wang Z, Li S, Zhou Y, Zhang Q, Xiao J. 2024. A cooking and eating quality evaluating system for whole grain black rice. Molecular Breeding, 45, 7.

Tian Y S, Fu X Y, Yang Z Q, Wang B, Gao J J, Wang M Q, Xu J, Han H J, Li Z J, Yao Q H, Peng R H. 2020. Metabolic engineering of rice endosperm for betanin biosynthesis. New Phytologist, 225, 1915-1922.

Tian Y S, Xu J, Wang B, Fu X Y, Gao J J, Han H J, Li Z J, Wang L J, Zhang F J, Zhang W H, Deng Y D, Wang Y, Peng R H, Yao Q H. 2021. Riboflavin fortification of rice endosperm by metabolic engineering. Plant Biotechnology Journal, 19, 1483-1485.

Tokuhara D, Álvarez B, Mejima M, Hiroiwa T, Takahashi Y, Kurokawa S, Kuroda M, Oyama M, Kozuka-Hata H, Nochi T, Sagara H, Aladin F, Marcotte H, Frenken L G J, Iturriza-Gómara M, Kiyono H, Hammarström L, Yuki Y. 2013. Rice-based oral antibody fragment prophylaxis and therapy against rotavirus infection. The Journal of Clinical Investigation, 123, 3829-3838.

Trijatmiko K R, Dueñas C, Tsakirpaloglou N, Torrizo L, Arines F M, Adeva C, Balindong J, Oliva N, Sapasap M V, Borrero J, Rey J, Francisco P, Nelson A, Nakanishi H, Lombi E, Tako E, Glahn R P, Stangoulis J, Chadha-Mohanty P, Johnson A A T, et al. 2016. Biofortified indica rice attains iron and zinc nutrition dietary targets in the field. Scientific Reports, 6, 19792.

Trkola A, Purtscher M, Muster T, Ballaun C, Buchacher A, Sullivan N, Srinivasan K, Sodroski J, Moore J P, Katinger H. 1996. Human monoclonal antibody 2G12 defines a distinctive neutralization epitope on the gp120 glycoprotein of human immunodeficiency virus type 1. Journal of Virology, 70, 1100-1108.

Tsuji N, Suzuki K, Kasuga-Aoki H, Isobe T, Arakawa T, Matsumoto Y. 2003. Mice intranasally immunized with a recombinant 16-kilodalton antigen from roundworm Ascaris parasites are protected against larval migration of Ascaris suum. Infection and Immunity, 71, 5314-5323.

Vamvaka E, Arcalis E, Ramessar K, Evans A, O'keefe B R, Shattock R J, Medina V, Stöger E, Christou P, Capell T. 2016a. Rice endosperm is cost-effective for the production of recombinant griffithsin with potent activity against HIV. Plant Biotechnology Journal, 14, 1427-1437.

Vamvaka E, Farré G, Molinos-Albert L M, Evans A, Canela-Xandri A, Twyman R M, Carrillo J, Ordóñez R A, Shattock R J, O’keefe B R, Clotet B, Blanco J, Khush G S, Christou P, Capell T. 2018. Unexpected synergistic HIV neutralization by a triple microbicide produced in rice endosperm. Proceedings of the National Academy of Sciences of the United States of America, 115, E7854-E7862.

Vamvaka E, Twyman R M, Murad A M, Melnik S, Teh A Y H, Arcalis E, Altmann F, Stoger E, Rech E, Ma J K C, Christou P, Capell T. 2016b. Rice endosperm produces an underglycosylated and potent form of the HIV-neutralizing monoclonal antibody 2G12. Plant Biotechnology Journal, 14, 97-108.

Vernet A, Meynard D, Lian Q, Mieulet D, Gibert O, Bissah M, Rivallan R, Autran D, Leblanc O, Meunier A C, Frouin J, Taillebois J, Shankle K, Khanday I, Mercier R, Sundaresan V, Guiderdoni E. 2022. High-frequency synthetic apomixis in hybrid rice. Nature Communications, 13, 7963.

Wakasa Y, Kawakatsu T, Ishimaru K, Ozawa K. 2024. Generation of major glutelin-deficient (GluA, GluB, and GluC) semi-dwarf Koshihikari rice line. Plant Cell Reports, 43, 51.

Wang A, Cheng Q, Li W, Kan M, Zhang Y, Meng X, Guo H, Jing Y, Chen M, Liu G, Wu D, Li J, Yu H. 2025. Creation of high-resistant starch rice through systematic editing of amylopectin biosynthetic genes in rs4. Plant Biotechnology Journal, 23, 480-488.

Wang A, Jing Y, Cheng Q, Zhou H, Wang L, Gong W, Kou L, Liu G, Meng X, Chen M, Ma H, Shu X, Yu H, Wu D, Li J. 2023. Loss of function of SSIIIa and SSIIIb coordinately confers high RS content in cooked rice. Proceedings of the National Academy of Sciences of the United States of America, 120, e2220622120.

Wang C, Liu Q, Shen Y, Hua Y, Wang J, Lin J, Wu M, Sun T, Cheng Z, Mercier R, Wang K. 2019. Clonal seeds from hybrid rice by simultaneous genome engineering of meiosis and fertilization genes. Nature Biotechnology, 37, 283-286.

Wang C, Wang K, Kou Y. 2024. Genome editing creates disease-resistant crops without yield penalties. Trends in Plant Science, 29, 114-116.

Wang J, Yang X, Fan L, Ye H, Hao G, Wang P. 2025a. The rational design of a Rhodamine fluorescent probe enables the selective detection and bioimaging of salicylic acid in plants under abiotic stress. Plant Biotechnology Journal, 23, 1736-1750.

Wang J, Zhang L, Wang S, Wang X, Li S, Gong P, Bai M, Paul A, Tvedt N, Ren H, Yang M, Zhang Z, Zhou S, Sun J, Wu X, Kuang H, Du Z, Dong Y, Shi X, Li M, et al. 2025b. AlphaFold-guided bespoke gene editing enhances field-grown soybean oil contents. Advanced Science, 12, 2500290.

Wang L M, Shen B R, Li B D, Zhang C L, Lin M, Tong P P, Cui L L, Zhang Z S, Peng X X. 2020. A synthetic photorespiratory shortcut enhances photosynthesis to boost biomass and grain yield in rice. Molecular Plant, 13, 1802-1815.

Wang T, Qin B R, Li S, Wang Z, Li X, Jiang Y, Qin C, Ouyang Q, Lou C, Qian L. 2025. Discovery of diverse and high-quality mRNA capping enzymes through a language model–enabled platform. Science Advances, 11, eadt0402.

Wang W, Mauleon R, Hu Z, Chebotarov D, Tai S, Wu Z, Li M, Zheng T, Fuentes R R, Zhang F, Mansueto L, Copetti D, Sanciangco M, Palis K C, Xu J, Sun C, Fu B, Zhang H, Gao Y, Zhao X, et al. 2018. Genomic variation in 3,010 diverse accessions of Asian cultivated rice. Nature, 557, 43-49.

Wang Y, Florez I D, Morgan R L, Foroutan F, Chang Y, Crandon H N, Zeraatkar D, Bala M M, Mao R Q, Tao B, Shahid S, Wang X, Beyene J, Offringa M, Sherman P M, El Gouhary E, Guyatt G H, Sadeghirad B. 2023. Probiotics, prebiotics, lactoferrin, and combination products for prevention of mortality and morbidity in preterm infants: A systematic review and network meta-analysis. JAMA Pediatrics, 177, 1158-1167.

Wang Y, Lilienfeldt N, Hekimi S. 2024. Understanding coenzyme Q. Physiological Reviews, 104, 1533-1610.

Wei X, Liu C, Chen X, Lu H, Wang J, Yang S, Wang K. 2023. Synthetic apomixis with normal hybrid rice seed production. Molecular Plant, 16, 489-492.

Wei Y, Zhang H, Fan J, Cai Q, Zhang Z, Wang J, Zhang M, Yan F, Jiang J, Xie H, Luo X, Wei L, Lin Y, He W, Qu M, Zhang X, Zhu Y, Xie H, Zhang J. 2024. Multiplex-genome-editing based rapid directional improvement of complex traits in rice. Plant Biotechnology Journal, 22, 2624-2628.

Williams M, Leemans J. 1993. Maintenance of Male-Sterile Plants. U.S. Patent, PCT Pub. No. WO93/ 25695. Patent and Trademark Office, Washington, D.C., USA.

Wu J, Qiu S, Wang M, Xu C, Deng X W, Tang X. 2021. Construction of a weight-based seed sorting system for the third-generation hybrid rice. Rice, 14, 66.

Wu S, Jin K, Li H, Chen G, Zhang L, Yang J, Zhai S, Li Y, Sun X, Cui X, Sun J, Lu T, Zhang Z. 2025. A synthetic facultative CAM-like shuttle in C3 rice plants. Advanced Science, 12, e2500418.

Wu W, Miao R, Li Z, Fang Z. 2025. CRISPR/Cas9-mediated editing of BADH2 and Wx genes for the development of novel aromatic and soft-textured black and red rice. Physiologia Plantarum, 177, e70194.

Wu Y, Fox T W, Trimnell M R, Wang L, Xu R J, Cigan A M, Huffman G A, Garnaat C W, Hershey H, Albertsen M C. 2016. Development of a novel recessive genetic male sterility system for hybrid seed production in maize and other cross-pollinating crops. Plant Biotechnology Journal, 14, 1046-1054.

Xia Y, Tang N, Hu Y, Li D, Li S, Bu X, Yu M, Qi S, Yang Y, Zhu H, Cao C, Li P, Yuan L, Cao M. 2019. A method for mechanized hybrid rice seed production using female sterile rice. Rice, 12, 39.

Xie E, Li Y, Tang D, Lv Y, Shen Y, Cheng Z. 2019. A strategy for generating rice apomixis by gene editing. Journal of Integrative Plant Biology, 61, 911-916.

Xie T, Qiu Q, Zhang W, Ning T, Yang W, Zheng C, Wang C, Zhu Y, Yang D. 2008. A biologically active rhIGF-1 fusion accumulated in transgenic rice seeds can reduce blood glucose in diabetic mice via oral delivery. Peptides, 29, 1862-1870.

Xie X, Moyu Z, Xinxin H, Xinrui Y, Nasser M, Ussemane M C, Ren M. 2025. Astaxanthin biosynthesis for functional food development and space missions. Critical Reviews in Biotechnology, 45, 923-937.

Xiong J, Ji Y, Yang S, Huang Y, Qiu X, Qian Q, Underwood C J, Wang K. 2025. Extending Mendel's legacy: The application of hawkweed PpPAR for inducing synthetic apomixis in hybrid rice. Plant Communications, 101387.

Xu H, Wang H, Zhang Y, Yang X, Lv S, Hou D, Mo C, Wassie M, Yu B, Hu T. 2023. A synthetic light-inducible photorespiratory bypass enhances photosynthesis to improve rice growth and grain yield. Plant Communications, 4, 100641.

Xu J J, Lei Y, Zhang X F, Li J X, Lin Q, Wu X D, Jiang Y G, Zhang W, Qian R, Xiong S Y, Tan K, Jia Y, Zhou Q, Jiang Y, Fan H, Huang Y B, Wang L J, Liu J Y, Kong Y, Zhao Q, et al. 2025. Design of CoQ10 crops based on evolutionary history. Cell, 188, 1941-1954.

Xu N, Yu D, Zhang L, Wang T, Hu Z, Lei Q, Qiu Y, Yuan Z, Song S, Li L. 2025. Establishment of a novel engineering breeding technology for cytoplasmic male sterility. The Crop Journal, 13, 1305-1311.

Xu Z, Xu X, Gong Q, Li Z, Li Y, Wang S, Yang Y, Ma W, Liu L, Zhu B, Zou L, Chen G. 2019. Engineering broad-spectrum bacterial blight resistance by simultaneously disrupting variable TALE-binding elements of multiple susceptibility genes in rice. Molecular Plant, 12, 1434-1446.

Yang F, Kong H, Ying J, Chen Z, Luo T, Jiang W, Yuan Z, Wang Z, Ma Z, Wang S, Ma W, Wang X, Li X, Hu Z, Ma X, Liu M, Wang X Q, Chen F, Dong N. 2025. SeedLLM·Rice: A large language model integrated with rice biological knowledge graph. Molecular Plant, 18, 1118-1129.

Yang Q, Tang X, Wu Y, Zhu W, Zhang T, Zhang Y. 2024. CRISPR-based modulation of uORFs in DEP1 and GIF1 for enhanced rice yield traits. Rice, 17, 67.

Yang R, Bai J, Fang J, Wang Y, Lee G, Piao Z. 2016. A single amino acid mutation of OsSBEIIb contributes to resistant starch accumulation in rice. Breeding Science, 66, 481-489.

Yang X, Wu D, Shi J, He Y, Pinot F, Grausem B, Yin C, Zhu L, Chen M, Luo Z, Liang W, Zhang D. 2014. Rice CYP703A3, a cytochrome P450 hydroxylase, is essential for development of anther cuticle and pollen exine. Journal of Integrative Plant Biology, 56, 979-994.

Yang Y, Shen Z, Li Y, Xu C, Xia H, Zhuang H, Sun S, Guo M, Yan C. 2022. Rapid improvement of rice eating and cooking quality through gene editing toward glutelin as target. Journal of Integrative Plant Biology, 64, 1860-1865.

Yang Z, Li D, Wei R, Zhao X, Lian X, Ren C, Xiao J, Li Y, Chen X, Wing R A, Zhang Q, Chen H. 2025. Whole grain black rice: A paradigm shift from mainly calories to a nutritious food that benefits human health. Science China Life Sciences, 68, 2173-2176.

Yao L, Zhang Y, Liu C, Liu Y, Wang Y, Liang D, Liu J, Sahoo G, Kelliher T. 2018. OsMATL mutation induces haploid seed formation in indica rice. Nature Plants, 4, 530-533.

Ye X, Al-Babili S, Klöti A, Zhang J, Lucca P, Beyer P, Potrykus I. 2000. Engineering the provitamin A (β-carotene) biosynthetic pathway into (carotenoid-free) rice endosperm. Science, 287, 303-305.

Ye X, Qin K, Fernie A R, Zhang Y. 2024. Prospects for synthetic biology in 21st Century agriculture. Journal of Genetics and Genomics, 52, 967-986.

Yi X, Li C. 2022. Main controllers for improving the resistant starch content in cooked white rice. Food Hydrocolloids, 122, 107083.

Yin T, Tai Y, Sun Y, Cheng Z, Wu C, Sui Y. 2025. Precise tiller angle control by manipulating TAC1 expression in rice. Journal of Integrative Plant Biology, 67, 1444-1446.

Yoshida T, Kimura E, Koike S, Nojima J, Futai E, Sasagawa N, Watanabe Y, Ishiura S. 2011. Transgenic rice expressing amyloid β-peptide for oral immunization. International Journal of Biological Sciences, 7, 301-307.

Yuan L P. 1987. Strategic conception of hybrid rice breeding. Hybrid Rice, 1, 1-3. (in Chinese)

Yuki Y, Mejima M, Kurokawa S, Hiroiwa T, Kong I G, Kuroda M, Takahashi Y, Nochi T, Tokuhara D, Kohda T, Kozaki S, Kiyono H. 2012. RNAi suppression of rice endogenous storage proteins enhances the production of rice-based Botulinum neutrotoxin type A vaccine. Vaccine, 30, 4160-4166.

Yuki Y, Nojima M, Hosono O, Tanaka H, Kimura Y, Satoh T, Imoto S, Uematsu S, Kurokawa S, Kashima K, Mejima M, Nakahashi-Ouchida R, Uchida Y, Marui T, Yoshikawa N, Nagamura F, Fujihashi K, Kiyono H. 2021. Oral MucoRice-CTB vaccine for safety and microbiota-dependent immunogenicity in humans: A phase 1 randomised trial. The Lancet Microbe, 2, e429-e440.

Zhan Y, Xia Y, Wang Y, Liu S, Zhang X, Xiong S, Lv Q, Cao M. 2025. Efficient clonal seeds sorting for apomictic hybrid rice using a pollen-specific gene switch system. Plant Biotechnology Journal, 23, 2266–2275.

Zhang L, Shi J, Jiang D, Stupak J, Ou J, Qiu Q, An N, Li J, Yang D. 2013. Expression and characterization of recombinant human alpha-antitrypsin in transgenic rice seed. Journal of Biotechnology, 164, 300-308.

Zhang L X, Li C X, Kakar M U, Khan M S, Wu P F, Amir R M, Dai D F, Naveed M, Li Q Y, Saeed M, Shen J Q, Rajput S A, Li J H. 2021. Resveratrol (RV): A pharmacological review and call for further research. Biomedicine & Pharmacotherapy, 143, 112164.

Zhang R, Wang Y, Yang W, Wen J, Liu W, Zhi S, Li G, Chai N, Huang J, Xie Y, Xie X, Chen L, Gu M, Liu Y G, Zhu Q. 2025. PlantGPT: An Arabidopsis-based intelligent agent that answers questions about plant functional genomics. Advanced Science, 12, e03926.

Zhang X. 2019. Synthetic biology in China: Review and prospects. Scientia Sinica (Vitae), 49, 1543-1572. (in Chinese)

Zhao G P. 2018. Synthetic biology: Unsealing the convergence era of life science research. Bulletin of Chinese Academy of Sciences, 33, 1135-1149. (in Chinese)

Zhao Y, Hu J, Zhou Z, Li L, Zhang X, He Y, Zhang C, Wang J, Hong G. 2024. Biofortified rice provides rich sakuranetin in endosperm. Rice, 17, 19.

Zheng X, Wei F, Cheng C, Qian Q. 2024. A historical review of hybrid rice breeding. Journal of Integrative Plant Biology, 66, 532-545.

Zhong Z, Fan T, He Y, Liu S, Zheng X, Xu Y, Ren J, Yuan H, Xu Z, Zhang Y. 2024. An improved plant prime editor for efficient generation of multiple-nucleotide variations and structural variations in rice. Plant Communications, 5, 100976.

Zhou H, Wang L, Liu G, Meng X, Jing Y, Shu X, Kong X, Sun J, Yu H, Smith S M, Wu D, Li J. 2016. Critical roles of soluble starch synthase SSIIIa and granule-bound starch synthase Waxy in synthesizing resistant starch in rice. Proceedings of the National Academy of Sciences of the United States of America, 113, 12844-12849.

Zhou S, Cai L, Wu H, Wang B, Gu B, Cui S, Huang X, Xu Z, Hao B, Hou H, Hu Y, Li C, Tian Y, Liu X, Chen L, Liu S, Jiang L, Wan J. 2024. Fine-tuning rice heading date through multiplex editing of the regulatory regions of key genes by CRISPR-Cas9. Plant Biotechnology Journal, 22, 751-758.

Zhu J, Zhou S, Wang W, Lv Y, Yang C, Wang Q, Zhang R, Shen S, Luo J. 2025. Development of “Tea Rice” by engineering catechin biosynthesis in rice endosperm. Plant Biotechnology Journal, 23, 2364-2366.

Zhu M, Yan L, Zhan Z, Chen H, Wang D, Xu M, Zheng Z, Zhang Y, Yang N, Wu J, Zhan H, Tian Y, Xiong L, He Y. 2025. Transgene-Killer-CRISPR version 2 (TKC2) eliminates occasional transgene escape by coupling with a RUBY reporter. Plant Biotechnology Journal, 23, 4621-4632.

Zhu Q, Tan J, Liu Y G. 2022. Molecular farming using transgenic rice endosperm. Trends in Biotechnology, 40, 1248-1260.

Zhu Q, Yu S, Zeng D, Liu H, Wang H, Yang Z, Xie X, Shen R, Tan J, Li H, Zhao X, Zhang Q, Chen Y, Guo J, Chen L, Liu Y G. 2017. Development of “Purple Endosperm Rice” by engineering anthocyanin biosynthesis in the endosperm with a high-efficiency transgene stacking system. Molecular Plant, 10, 918-929.

Zhu Q, Zeng D, Yu S, Cui C, Li J, Li H, Chen J, Zhang R, Zhao X, Chen L, Liu Y G. 2018. From golden rice to aSTARice: Bioengineering astaxanthin biosynthesis in rice endosperm. Molecular Plant, 11, 1440-1448.

No related articles found!
No Suggested Reading articles found!