The research report should be a detailed narrative explaining the function, biological processes, and localization of the gene product. Citations should be given for all claims.
You should prioritize authoritative reviews and primary scientific literature when conducting research. You can supplement
this with annotations you find in gene/protein databases, but these can be outdated or inaccurate.
We are specifically interested in the primary function of the gene - for enzymes, what reaction is catalyzed, and what is the substrate specificity? For transporters, what is the substrate? For structural proteins or adapters, what is the broader structural role? For signaling molecules, what is the role in the pathway.
We are interested in where in or outside the cell the gene product carries out its function.
We are also interested in the signaling or biochemical pathways in which the gene functions. We are less interested in broad pleiotropic effects, except where these elucidate the precise role.
Include evidence where possible. We are interested in both experimental evidence as well as inference from structure, evolution, or bioinformatic analysis. Precise studies should be prioritized over high-throughput, where available.
CKX2 (OsCKX2; Gn1a) encodes a cytokinin oxidase/dehydrogenase that irreversibly degrades cytokinins, thereby controlling cytokinin homeostasis in inflorescence meristems and developing panicles and tuning meristem activity that ultimately determines panicle branching and grain number in rice. Genetic reduction or loss of OsCKX2 activity increases cytokinin accumulation in reproductive meristems and can substantially increase grain number and yield components, making OsCKX2 one of the most validated hormone-metabolism targets for rice yield improvement. (ashikari2005cytokininoxidaseregulates pages 1-2, yeh2015downregulationofcytokinin pages 1-2)
Cytokinins are adenine-derived phytohormones that regulate plant growth and development, including meristem activity. Their steady-state levels depend on biosynthesis, interconversion among nucleobase/nucleoside/nucleotide forms, conjugation, transport, and irreversible degradation. Cytokinin oxidase/dehydrogenase (CKX) enzymes mediate the key irreversible catabolic step by cleaving cytokinin side chains. (werner12006newinsightsinto pages 1-2)
CKX enzymes catalyze irreversible cytokinin degradation by cleaving the unsaturated isoprenoid N6 side chain of cytokinin substrates to yield adenine/adenosine plus a corresponding aldehyde side-chain product. (werner12006newinsightsinto pages 1-2, werner12006newinsightsinto pages 6-7)
At the mechanistic level, CKXs are FAD-dependent flavoenzymes. The cytokinin substrate reacts with oxidized FAD through a transient adduct, leading to formation of an intermediate that is hydrolyzed to adenine and an aldehyde; CKX can operate in an “oxidase” mode (using oxygen weakly) or more efficiently as a “dehydrogenase” using electron acceptors such as quinones, consistent with the dehydrogenase nomenclature. (werner12006newinsightsinto pages 6-7)
The target protein is UniProt Q4ADV8 from Oryza sativa subsp. japonica (rice), annotated as cytokinin dehydrogenase/oxidase 2 with aliases OsCKX2 and Gn1a. In rice genetics, Gn1a is a grain-number QTL on chromosome 1 that was molecularly cloned and shown to correspond to OsCKX2 (cytokinin oxidase/dehydrogenase). (ashikari2005cytokininoxidaseregulates pages 1-2, ashikari2005cytokininoxidaseregulates pages 2-3)
High-resolution mapping and transgenic confirmation established that the rice QTL Gn1a corresponds to OsCKX2, a predicted cytokinin oxidase/dehydrogenase gene in rice (Oryza sativa). This was shown in the context of japonica cultivar Koshihikari and the high-yield indica cultivar Habataki, among other materials. (ashikari2005cytokininoxidaseregulates pages 1-2, ashikari2005cytokininoxidaseregulates pages 2-3)
A figure set in the original cloning paper shows the genetic mapping of Gn1a to OsCKX2 and accompanying cytokinin/OsCKX2 expression data in inflorescence meristems. (ashikari2005cytokininoxidaseregulates media 9c2fb589)
OsCKX2 is a CKX-family cytokinin catabolic enzyme whose reduced activity leads to accumulation of cytokinin species (including nucleotide forms such as tZRMP and iPRMP) in inflorescence meristems, consistent with decreased cytokinin degradation. OsCKX2 protein variants expressed in yeast retain cytokinin-cleaving activity, supporting that OsCKX2 encodes an active CKX enzyme. (ashikari2005cytokininoxidaseregulates pages 3-4)
Family-level enzymology indicates CKXs have highest affinity for isoprenoid cytokinins such as isopentenyladenine (iP) and trans-zeatin (tZ) (and their ribosides), often with low-micromolar Km values. (werner12006newinsightsinto pages 6-7)
OsCKX2-specific substrate evidence available in the retrieved corpus is primarily functional (in planta) rather than kinetic/enzymatic, but knockdown/reduced-expression OsCKX2 backgrounds are associated with elevated active cytokinin forms including tZ and iP-type cytokinins in reproductive meristems, consistent with OsCKX2 acting on those substrates. (ashikari2005cytokininoxidaseregulates pages 1-2, katerUnknownyear…cytokininmetabolism pages 17-19)
OsCKX2 is a negative regulator of cytokinin levels in inflorescence meristems/young panicles, and thus acts as a negative regulator of meristem activity and reproductive organ production. Reduced OsCKX2 expression leads to cytokinin accumulation in inflorescence meristems and increases the number of reproductive organs, increasing grain yield. (ashikari2005cytokininoxidaseregulates pages 1-2, ashikari2005cytokininoxidaseregulates pages 3-4)
In the molecular cloning study of Gn1a, the Habataki Gn1a allele was reported to be semidominant and associated with ~92 more grains per main panicle, accounting for 44% of the grain-number difference relative to Koshihikari. A natural null allele (11-bp deletion) in variety 5150 was associated with >400 grains per main panicle, consistent with the concept that reduced or lost OsCKX2 function enhances grain production. (ashikari2005cytokininoxidaseregulates pages 2-3)
A field-evaluated shRNA knockdown strategy targeting OsCKX2 in rice demonstrated that OsCKX2 suppression increases multiple yield components. In field experiments, transgenic lines produced 27–81% more tillers and 24–67% more grains per plant, and showed 5–15% heavier 1000-grain weight than wild-type plants. The authors also report that insertional activation (increased expression) of OsCKX2 reduces tiller number and growth in a gene-dosage–dependent manner. (yeh2015downregulationofcytokinin pages 1-2)
A 2023 study (BMC Plant Biology; publication date Dec 2023; DOI URL https://doi.org/10.1186/s12870-023-04671-4) provides mechanistic insight into how OsCKX2 is transcriptionally positioned in panicle/spikelet development.
The study shows that FZP, a nuclear AP2/ERF transcription factor, represses DST by directly binding its promoter; in a severe fzp allele, DST and OsCKX2 are upregulated in young panicles, and cytokinin content decreases substantially (IP ~60% lower, IPA ~25% lower, zeatin ~27% lower; DHZ unchanged). The severe fzp phenotype included a ~66% reduction in grain number per main panicle, linking an upstream developmental regulator to OsCKX2-mediated cytokinin depletion and reduced grain number. (wang2023frizzlepanicle(fzp) pages 2-4, wang2023frizzlepanicle(fzp) pages 1-2)
In addition, the same work synthesizes a regulatory concept in which DST binds the OsCKX2 promoter and its activity can be enhanced by phosphorylation via the OsMKKK10–OsMKK4–OsMPK6 cascade; MED25 is described as a DST coactivator, situating OsCKX2 in a broader meristem-patterning signaling context. (wang2023frizzlepanicle(fzp) pages 1-2)
A 2024 study (Plant Cell Reports; publication date Aug 2024; DOI URL https://doi.org/10.1007/s00299-024-03289-6) generated OsCKX2-deficient mutants using CRISPR/Cas9 in indica rice (MTU1010). The study reports that Osckx2 loss-of-function increases cytokinin accumulation in panicle tissue and improves panicle architecture (including increased secondary branching), grain number, and overall yield, and additionally enhances drought-related performance (reduced transpiration, improved survival under dehydration stress, better membrane/chloroplast integrity, and improved photosynthetic function through antioxidant protection). Within the excerpted pages available here, numeric effect sizes were not provided, but the mechanistic interpretation is consistent with OsCKX2 functioning as a negative regulator of reproductive cytokinin levels and panicle sink strength. (rashid2024cytokininoxidase2deficientmutants pages 1-5, rashid2024cytokininoxidase2deficientmutants pages 5-7)
Direct subcellular localization evidence for OsCKX2 protein (e.g., ER lumen vs apoplast) was not present in the retrieved full-text snippets. However, multiple authoritative sources emphasize that CKX family proteins have diverse subcellular targeting (extracellular/apoplastic, vacuolar, cytosolic, ER-associated) and that localization differences are an important axis of CKX functional specialization. In rice-family examples, OsCKX3 is described as ER-localized (GFP fusion), while OsCKX4 was reported as cytosolic, illustrating that rice CKX paralogs are not uniformly localized. (werner12006newinsightsinto pages 2-3, katerUnknownyear…cytokininmetabolism pages 17-19)
Functionally, OsCKX2 action is repeatedly localized at the tissue level to inflorescence meristems/young panicles, where changes in OsCKX2 expression are coupled to changes in cytokinin levels and panicle/grain phenotypes. (ashikari2005cytokininoxidaseregulates pages 1-2, wang2023frizzlepanicle(fzp) pages 2-4)
OsCKX2/Gn1a is a canonical example of a yield QTL molecularly traced to a hormone catabolic enzyme. The large effect sizes reported for the Habataki allele (e.g., +92 grains per main panicle; 44% of difference vs Koshihikari) provide a rationale for marker-assisted introgression and QTL pyramiding strategies that incorporate sink-size loci. (ashikari2005cytokininoxidaseregulates pages 2-3, ashikari2005cytokininoxidaseregulates pages 1-2)
The shRNA knockdown study demonstrates a path from molecular perturbation to field performance, showing that OsCKX2 suppression can increase tiller number, grains per plant, and 1000-grain weight in field conditions. (yeh2015downregulationofcytokinin pages 1-2)
The 2024 indica-rice CRISPR/Cas9 study illustrates OsCKX2 as a target for precision breeding aimed at simultaneously improving sink traits (panicle branching, grain number) and climate resilience (drought tolerance traits). (rashid2024cytokininoxidase2deficientmutants pages 1-5, rashid2024cytokininoxidase2deficientmutants pages 5-7)
A consistent expert consensus across primary studies and mechanistic reviews is that cytokinin degradation is a leverage point for yield manipulation because cytokinins promote meristem activity and reproductive organ production. OsCKX2/Gn1a exemplifies this by connecting a defined enzymatic step (cytokinin side-chain cleavage) to a high-impact agronomic trait (grain number). (werner12006newinsightsinto pages 6-7, ashikari2005cytokininoxidaseregulates pages 1-2)
Mechanistically, the 2023 FZP–DST–OsCKX2 study suggests that yield effects mediated through OsCKX2 are not simply “more cytokinin is better”; rather, OsCKX2 must be tuned within a regulatory network balancing meristem fate, spikelet development, and cytokinin levels, because excessive cytokinin depletion (via upregulated OsCKX2) can reduce grain number dramatically (e.g., ~66% reduction in the severe fzp allele context). (wang2023frizzlepanicle(fzp) pages 1-2)
| Topic | Key points | Quantitative data | Organism/background | Supporting citation IDs |
|---|---|---|---|---|
| Identity/locus | OsCKX2 is the rice cytokinin oxidase/dehydrogenase 2 gene; the grain-number QTL Gn1a was molecularly cloned as OsCKX2 on chromosome 1. Reduced-expression or null alleles increase cytokinin in inflorescence meristems and raise grain number. This matches the UniProt target CKX2/OsCKX2/Gn1a from Oryza sativa. | Habataki Gn1a allele increased grain number by ~92 grains per main panicle and explained 44% of the grain-number difference versus Koshihikari; a natural 11-bp deletion allele in 5150 was associated with >400 grains per main panicle. | Rice (Oryza sativa), especially japonica Koshihikari and indica Habataki/5150 backgrounds | (ashikari2005cytokininoxidaseregulates pages 2-3, ashikari2005cytokininoxidaseregulates pages 1-2) |
| Enzymatic function | OsCKX2 encodes a CKX-family enzyme that irreversibly degrades cytokinins, thereby lowering endogenous CK abundance in reproductive meristems and other tissues. Functional expression in yeast confirmed CKX catalytic activity for OsCKX2 protein variants. | Reduced OsCKX2 activity in Gn1a-associated lines caused accumulation of cytokinin nucleotides such as tZRMP and iPRMP relative to Koshihikari. | Rice Gn1a/OsCKX2 lines; yeast heterologous expression for catalytic confirmation | (ashikari2005cytokininoxidaseregulates pages 3-4, yeh2015downregulationofcytokinin pages 1-2) |
| Substrates & mechanism | Plant CKX enzymes are FAD-dependent flavoenzymes that cleave the unsaturated N6 isoprenoid side chain of cytokinin nucleobases/ribosides, yielding adenine/adenosine plus an aldehyde. Family-level evidence indicates preference for isoprenoid CKs such as iP and tZ (and their ribosides), supporting inference for OsCKX2. Rice-specific knockdown evidence indicates OsCKX2 normally restrains tZ, 2-iP, kinetin, and DHZ in inflorescence meristems. | CKX enzymes have low-micromolar Km for iP/tZ-type substrates in family-level studies; OsCKX2-RNAi/knockdown plants showed elevated tZ, 2-iP, kinetin, DHZ in IM tissue. | General plant CKX enzymology with rice OsCKX2 functional inference; rice inflorescence meristem knockdown lines | (werner12006newinsightsinto pages 6-7, werner12006newinsightsinto pages 1-2, katerUnknownyear…cytokininmetabolism pages 17-19, katerUnknownyear…cytokininmetabolisma pages 17-19) |
| Localization | Direct OsCKX2 localization evidence was not retrieved in the gathered texts. However, OsCKX2 has a signal peptide in UniProt and CKX family proteins localize to distinct compartments including extracellular/apoplastic, ER-associated, vacuolar, and cytosolic pools depending on isoform. OsCKX2 function is experimentally tied to the inflorescence meristem/panicle tissue where its transcript abundance affects CK levels. | No direct OsCKX2 compartment quantification retrieved; OsCKX3 localizes predominantly to ER, OsCKX4 is cytosolic, illustrating family divergence. | Rice CKX family context; OsCKX2 expression/function studied in inflorescence meristems and young panicles | (werner12006newinsightsinto pages 2-3, katerUnknownyear…cytokininmetabolism pages 17-19, ashikari2005cytokininoxidaseregulates media 9c2fb589) |
| Pathway/regulation | OsCKX2 is a core node in cytokinin-homeostasis pathways controlling inflorescence meristem activity. DST directly activates OsCKX2; MED25 acts as a DST coactivator; phosphorylation by the OsMKKK10–OsMKK4–OsMPK6 cascade promotes DST-mediated OsCKX2 expression. FZP represses DST, thereby indirectly limiting OsCKX2 and sustaining CK in young panicles. | In the severe fzp/abp1 allele, grain number per main panicle fell ~66%; in fzp young panicles, CKs decreased: IP ~60% lower, IPA ~25% lower, zeatin ~27% lower. | Rice young panicles/inflorescence meristems; FZP-DST-OsCKX2 regulatory module | (wang2023frizzlepanicle(fzp) pages 1-2, wang2023frizzlepanicle(fzp) pages 2-4) |
| Phenotypes/yield data | Lower OsCKX2 expression increases sink strength by enhancing meristem activity, panicle branching, tillering, and grain number; overexpression/activation has the opposite effect. OsCKX2 is therefore a negative regulator of grain number and tiller formation through cytokinin catabolism. | shRNA suppression caused 27–81% more tillers, 24–67% more grains per plant, and 5–15% higher 1000-grain weight in field tests; insertional activation reduced tiller number in a dosage-dependent manner. | Transgenic rice, including O. sativa cv. TNG67; greenhouse and field experiments | (yeh2015downregulationofcytokinin pages 1-2, yeh2015downregulationofcytokinin pages 8-10) |
| Recent 2023–2024 developments | 2023 work refined the upstream transcriptional network controlling OsCKX2 via FZP→DST→OsCKX2 and linked altered OsCKX2 expression to reduced cytokinin and defective spikelet development. 2024 CRISPR work showed that Osckx2 loss-of-function in indica rice increases panicle branching and grain yield while also improving drought-related performance. | 2023: 66% reduction in grain number per main panicle in severe fzp background; CK decreases of ~60% IP, ~25% IPA, ~27% zeatin. 2024: qualitative increases in secondary branching, grain number, and drought survival traits; provided pages did not include exact numeric effect sizes. | 2023 BMC Plant Biology study in rice panicles; 2024 CRISPR/Cas9 study in indica rice cv. MTU1010 | (wang2023frizzlepanicle(fzp) pages 2-4, wang2023frizzlepanicle(fzp) pages 1-2, rashid2024cytokininoxidase2deficientmutants pages 1-5, rashid2024cytokininoxidase2deficientmutants pages 5-7) |
| Applications | OsCKX2/Gn1a is a validated target for yield breeding, QTL introgression, gene pyramiding, RNAi knockdown, and CRISPR/Cas9 editing. Practical use cases include exploiting weak/null alleles to raise grain number and using edited alleles for high-yield, climate-resilient rice. | Gn1a introgression delivered major sink-size effects (e.g., ~92 grains/panicle, 44% difference component); recent CRISPR mutants increased grain number and drought tolerance qualitatively; OsCKX2 knockdown also reduced yield penalty under salinity in earlier work. | Rice breeding in japonica and indica backgrounds; near-isogenic lines, transgenics, and CRISPR-edited plants | (ashikari2005cytokininoxidaseregulates pages 2-3, rashid2024cytokininoxidase2deficientmutants pages 1-5, ashikari2005cytokininoxidaseregulates pages 1-2) |
Table: This table compiles the main functional-annotation evidence for rice OsCKX2/Gn1a, including identity, enzymatic role, regulatory pathway, quantitative phenotypes, and 2023–2024 updates. It is useful as a compact evidence map for gene-function interpretation and breeding relevance.
References
(ashikari2005cytokininoxidaseregulates pages 1-2): Motoyuki Ashikari, Hitoshi Sakakibara, Shaoyang Lin, Toshio Yamamoto, Tomonori Takashi, Asuka Nishimura, Enrique R. Angeles, Qian Qian, Hidemi Kitano, and Makoto Matsuoka. Cytokinin oxidase regulates rice grain production. Science, 309:741-745, Jul 2005. URL: https://doi.org/10.1126/science.1113373, doi:10.1126/science.1113373. This article has 2477 citations and is from a highest quality peer-reviewed journal.
(yeh2015downregulationofcytokinin pages 1-2): Su-Ying Yeh, Hau-Wen Chen, Chun-Yeung Ng, Chu-Yin Lin, Tung-Hai Tseng, Wen-Hsiung Li, and Maurice S. B. Ku. Down-regulation of cytokinin oxidase 2 expression increases tiller number and improves rice yield. Rice, Dec 2015. URL: https://doi.org/10.1186/s12284-015-0070-5, doi:10.1186/s12284-015-0070-5. This article has 192 citations and is from a peer-reviewed journal.
(werner12006newinsightsinto pages 1-2): T. Werner1, I. Köllmer1, I. Bartrina1, K. Holst1, and T. Schmülling1. New insights into the biology of cytokinin degradation. Plant Biology, 8:371-381, May 2006. URL: https://doi.org/10.1055/s-2006-923928, doi:10.1055/s-2006-923928. This article has 398 citations and is from a peer-reviewed journal.
(werner12006newinsightsinto pages 6-7): T. Werner1, I. Köllmer1, I. Bartrina1, K. Holst1, and T. Schmülling1. New insights into the biology of cytokinin degradation. Plant Biology, 8:371-381, May 2006. URL: https://doi.org/10.1055/s-2006-923928, doi:10.1055/s-2006-923928. This article has 398 citations and is from a peer-reviewed journal.
(ashikari2005cytokininoxidaseregulates pages 2-3): Motoyuki Ashikari, Hitoshi Sakakibara, Shaoyang Lin, Toshio Yamamoto, Tomonori Takashi, Asuka Nishimura, Enrique R. Angeles, Qian Qian, Hidemi Kitano, and Makoto Matsuoka. Cytokinin oxidase regulates rice grain production. Science, 309:741-745, Jul 2005. URL: https://doi.org/10.1126/science.1113373, doi:10.1126/science.1113373. This article has 2477 citations and is from a highest quality peer-reviewed journal.
(ashikari2005cytokininoxidaseregulates media 9c2fb589): Motoyuki Ashikari, Hitoshi Sakakibara, Shaoyang Lin, Toshio Yamamoto, Tomonori Takashi, Asuka Nishimura, Enrique R. Angeles, Qian Qian, Hidemi Kitano, and Makoto Matsuoka. Cytokinin oxidase regulates rice grain production. Science, 309:741-745, Jul 2005. URL: https://doi.org/10.1126/science.1113373, doi:10.1126/science.1113373. This article has 2477 citations and is from a highest quality peer-reviewed journal.
(ashikari2005cytokininoxidaseregulates pages 3-4): Motoyuki Ashikari, Hitoshi Sakakibara, Shaoyang Lin, Toshio Yamamoto, Tomonori Takashi, Asuka Nishimura, Enrique R. Angeles, Qian Qian, Hidemi Kitano, and Makoto Matsuoka. Cytokinin oxidase regulates rice grain production. Science, 309:741-745, Jul 2005. URL: https://doi.org/10.1126/science.1113373, doi:10.1126/science.1113373. This article has 2477 citations and is from a highest quality peer-reviewed journal.
(katerUnknownyear…cytokininmetabolism pages 17-19): M KATER. … cytokinin metabolism during inflorescence meristem development to promote higher rice productivity through changes …. Unknown journal, Unknown year.
(wang2023frizzlepanicle(fzp) pages 2-4): W. Wang, Wenqiang Chen, Qinglong Liu, and Junmin Wang. Frizzle panicle (fzp) regulates rice spikelets development through modulating cytokinin metabolism. BMC Plant Biology, Dec 2023. URL: https://doi.org/10.1186/s12870-023-04671-4, doi:10.1186/s12870-023-04671-4. This article has 9 citations and is from a peer-reviewed journal.
(wang2023frizzlepanicle(fzp) pages 1-2): W. Wang, Wenqiang Chen, Qinglong Liu, and Junmin Wang. Frizzle panicle (fzp) regulates rice spikelets development through modulating cytokinin metabolism. BMC Plant Biology, Dec 2023. URL: https://doi.org/10.1186/s12870-023-04671-4, doi:10.1186/s12870-023-04671-4. This article has 9 citations and is from a peer-reviewed journal.
(rashid2024cytokininoxidase2deficientmutants pages 1-5): Afreen Rashid, V. Mohan M. Achary, M. Z. Abdin, Sangeetha Karippadakam, Hemangini Parmar, Varakumar Panditi, Ganesan Prakash, Pooja Bhatnagar-Mathur, and Malireddy K. Reddy. Cytokinin oxidase2-deficient mutants improve panicle and grain architecture through cytokinin accumulation and enhance drought tolerance in indica rice. Plant cell reports, 43 8:207, Aug 2024. URL: https://doi.org/10.1007/s00299-024-03289-6, doi:10.1007/s00299-024-03289-6. This article has 25 citations and is from a peer-reviewed journal.
(rashid2024cytokininoxidase2deficientmutants pages 5-7): Afreen Rashid, V. Mohan M. Achary, M. Z. Abdin, Sangeetha Karippadakam, Hemangini Parmar, Varakumar Panditi, Ganesan Prakash, Pooja Bhatnagar-Mathur, and Malireddy K. Reddy. Cytokinin oxidase2-deficient mutants improve panicle and grain architecture through cytokinin accumulation and enhance drought tolerance in indica rice. Plant cell reports, 43 8:207, Aug 2024. URL: https://doi.org/10.1007/s00299-024-03289-6, doi:10.1007/s00299-024-03289-6. This article has 25 citations and is from a peer-reviewed journal.
(werner12006newinsightsinto pages 2-3): T. Werner1, I. Köllmer1, I. Bartrina1, K. Holst1, and T. Schmülling1. New insights into the biology of cytokinin degradation. Plant Biology, 8:371-381, May 2006. URL: https://doi.org/10.1055/s-2006-923928, doi:10.1055/s-2006-923928. This article has 398 citations and is from a peer-reviewed journal.
(katerUnknownyear…cytokininmetabolisma pages 17-19): M KATER. … cytokinin metabolism during inflorescence meristem development to promote higher rice productivity through changes …. Unknown journal, Unknown year.
(yeh2015downregulationofcytokinin pages 8-10): Su-Ying Yeh, Hau-Wen Chen, Chun-Yeung Ng, Chu-Yin Lin, Tung-Hai Tseng, Wen-Hsiung Li, and Maurice S. B. Ku. Down-regulation of cytokinin oxidase 2 expression increases tiller number and improves rice yield. Rice, Dec 2015. URL: https://doi.org/10.1186/s12284-015-0070-5, doi:10.1186/s12284-015-0070-5. This article has 192 citations and is from a peer-reviewed journal.