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Cold-induced peptide signalling secures pollen resilience and crop yield

Nature (2026) Cite this article

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Abstract

Cold weather cause severe crop losses. Climate change exacerbates the unpredictability and frequency of such weather events, highlighting the need for cold-resilient crops1. Cold-induced pollen abortion and reproductive failure during flowering are major causes of yield losses2,3, yet the molecular mechanisms and signalling pathways that underlie cold resilience in pollen development remain unknown. Here we identify a subset of cold-responsive small signalling peptides in the RGF–GLV–CLEL family, SlRGF9 and SlRGF10, that control cold resilience in tomato pollen. After loss of function of SlRGF9 and SlRGF10, tomato plants (Solanum lycopersicum) showed no defects under normal conditions, but pollen abortion was observed after cold stress. The leucine-rich repeat receptor-like kinases, SlRGFR6 and SlSERK proteins, form cell-surface receptor complexes that bind to these cold-induced SlRGFs. Furthermore, SlRGF–SlRGFR6 signalling activates calcium influx through cyclic-nucleotide-gated channels, counteracting cold-delayed programmed cell death and ensuring tapetum degradation to support microspore development. Upregulating SlRGF9 and SlRGF10 in tomato plants prevents cold-induced yield losses by up to 52%. This cold-responsive peptide signalling pathway is conserved across dicots and monocots. For example, upregulation of RGF homologues in rice (Oryza sativa) boosts cold resilience in pollen and recovers 18.3% of grain yield loss. Our findings uncover a core peptide signalling axis that governs cold resilience in pollen and has broad potential for safeguarding crop productivity against cold stress.

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Fig. 1: Chilling-induced SlRGFs promote cold resilience in tomato pollen.
The alternative text for this image may have been generated using AI.
Fig. 2: Chilling-induced SlRGF9/10 functions occur through SlRGFR6.
The alternative text for this image may have been generated using AI.
Fig. 3: SlRGFR6 binds directly to SlRGF9/10 peptides and forms a cell-surface receptor complex with SlSERKs to trigger calcium influx.
The alternative text for this image may have been generated using AI.
Fig. 4: SlRGF–SlRGFR6 signalling triggers calcium influx through SlCNGC16/18.
The alternative text for this image may have been generated using AI.
Fig. 5: Moderate upregulation of SlRGF genes rescues chilling-induced yield loss in tomato and rice.
The alternative text for this image may have been generated using AI.

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Data availability

Gene and protein sequences used in this study were collected from the Sol Genomics Network (tomato; https://solgenomics.net/), Rice Genome Annotation Project (rice; https://rice.uga.edu/), TAIR10 (Arabidopsis; https://www.arabidopsis.org/) and The Plant Genomics Resource (https://phytozome-next.jgi.doe.gov/). Expression pattern analysis was conducted with published RNA-sequencing data (GEO: GSE40854; SRA: PRJNA631095). Uncropped gel and blot images are provided in Supplementary Fig. 4. Primers and annotated peptides are provided in Supplementary Tables 1–3. Source data are provided with this paper.

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Acknowledgements

We thank Y. Guo, P. Leng, Y. Liu and J. Chai for sharing vectors; W. Guo, Y. Yu and J. Chai for sharing cell line transfection methods; G. Lu and C. Pan for sharing RNA sequencing data; J. Li and M. Chen for providing the rice seeds; S. Zhou and C. Shi for handling the rice management; Y. Tian for technical assistance; Y. Yu for protoplast isolation; J. Mou for detection of radioactivity; Y. Ming for optimization of protein extraction methods; and all the other members in the lab for assay assistance and project discussion.

Funding

This study was financially supported by Natural Science Foundation of China grants 32388101 and 32225045 (to C.X.) and 32270427 and 32470431 (to J.C. and P.X.), a China Agricultural University Fund grant 2025RC042 (to W.S.) and a State Key Laboratory of Plant Environmental Resilience Fund grant SKLPERKF2502 (to W.S.).

Author information

Authors and Affiliations

  1. State Key Laboratory of Seed Innovation, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing, China

    Shudong Chen, Yupan Zou, Huanshuo Cui, Qingfeng Dong, Dandan Yang, Xiaozhen Huang, Shujing Cheng, Peiyong Xin, Jinfang Chu & Cao Xu

  2. College of Advanced Agricultural Sciences, University of Chinese Academy of Sciences, Beijing, China

    Shudong Chen, Huanshuo Cui, Qingfeng Dong, Jinfang Chu & Cao Xu

  3. CAS–JIC Centre of Excellence for Plant and Microbial Science (CEPAMS), Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing, China

    Shudong Chen, Yupan Zou, Huanshuo Cui, Qingfeng Dong, Dandan Yang, Xiaozhen Huang, Shujing Cheng, Peiyong Xin, Jinfang Chu & Cao Xu

  4. State Key Laboratory of Plant Environmental Resilience, College of Biological Sciences, China Agricultural University, Beijing, China

    Wen Song

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Contributions

C.X. designed and supervised the research. W.S. contributed critical ideas and guided the research. S. Chen, Q.D. and H.C. generated mutants and performed phenotyping experiments and yield trials. S. Chen performed TUNEL staining, RT-qPCR, ITC assay and organized the panels. Y.Z. performed the bioinformatic analysis. D.Y. contributed to the cell biology experiments. X.H. generated the Slserk mutants. S. Cheng, P.X. and J.C. analysed the sulfated peptides. C.X. wrote the manuscript with input from S. Chen and W.S. All authors read and agreed to the submitted manuscript.

Corresponding authors

Correspondence to Wen Song or Cao Xu.

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Competing interests

C.X., S. Chen, S. Cheng, P.X. and J.C. based at the Institute of Genetics and Developmental Biology, Chinese Academy of Sciences have submitted patent applications covering the gene application (PCT/CN2026/092697) and method of peptide detection (CN202610197060.X) reported in this study. The other authors declare no competing interests. The authors affirm that this disclosure does not alter their adherence to the journal’s policies on data sharing and publication.

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Extended data figures and tables

Extended Data Fig. 1 Chilling stress causes pollen abortion and yield loss in tomato.

a, Daytime and nighttime temperature medians over a course of 15 d during flower in solar greenhouse related to Fig. 1a–b, Extended Data Fig. 1b–c. b–c, Fresh fruit yield per plant (b) and quantification (c) at normal and chilling conditions. d–e, Comparison of fruit setting rate for self-pollination with manual pollination (using pollens from optimum growth conditions) under various low growth temperatures. Red arrowheads indicate flowers that successfully developed into fruits, while white arrowheads indicate aborted flowers. Yellow circles indicate aborted flowers at the end of the inflorescence. f, Flower size and developmental stages corresponding to pollen development. g, Workflow for tomato chilling treatment and recovery growth for assess fruit set and pollen viability. h–i, Alexander staining of pollen viability (h) and quantification (i) in wild-type under different temperature treatments. Yellow arrowheads: nonviable pollens. j, Schematic summarizing fruit set and pollen viability under different low temperatures. Scale bars are 2 cm (b, d), 50 μm (h).“N”: individual plant (c, e), inflorescence (i); data: box plots: median (central line); 25th and 75th percentiles (bounds); minima and maxima (whiskers) (c), mean ± SD (e, i); P-values: Mann-Whitney U test (c, e) and Kruskal-Wallis test (i). Schematics in f, g and j created in BioRender; Chen, S. https://biorender.com/uc0y98j and https://biorender.com/8duox87 (2026).

Source data

Extended Data Fig. 2 Discovery of SlRGF genes and phenotypes of tomato Slrgf9/10 null mutants.

a, Workflow for bioinformatic screening of putative SSPs. b, Distribution of putative small signaling peptide genes on tomato chromosomes. Gene density distributions for small peptide-encoding loci are represented by color-coded vertical bars. c, Expression heatmap of SSPs across tomato tissues. EVM/MVM/LVM/TM/FM/SIM/SYM indicate the early vegetative/middle vegetative/late vegetative/transitional/floral/sympodial inflorescence/sympodial shoot meristem, respectively. d, Differential-expression analysis of SSP genes in anthers under normal temperature vs. chilling. e, Expression heatmap of SlRGFs across tissues under normal temperature. f, RT–qPCR validation of SlRGF1/6 expression in different flower tissues. g, Time-course expression of SlRGFs under chilling. h, In silico identification of chilling-responsive cis-elements in the SlRGF9/10 promoters (PlantCARE). i–j, Quantification of signal intensity of SlRGF9/10pro: GUS transgenic lines in anthers under low temperature. Representative images of GUS and GFP fluorescence signals from the same tomato plant. k, CRISPR-Cas9 gRNA design and allelic information for Slrgf9/10 mutants. l–n, Representative phenotypes (l) and quantification of flowering time (m), flower number per inflorescence (n) of wild-type and Slrgf9/10 mutants grown at normal temperature. White asterisks: primary leaves; red arrowheads: primary inflorescences. o–p, Phenotypes (o) and fruit setting rate quantification (p) of wild-type, Slrgf9, Slrgf10, and Slrgf9 Slrgf10 double mutants under normal temperature and chilling treatment. Red and white arrowheads indicate flowers at tetrads and early uninucleate stages subject to chilling treatment that successfully developed into fruits and aborted flowers, respectively. q, Anther section of SlRGFpro: GFP transgenic lines. r, Semi-thin anther sections of wild-type and Slrgf9 Slrgf10 mutants grown under normal temperature. s, TUNEL assay of tapetum PCD for wild-type and Slrgf9 Slrgf10 mutants grown under normal temperature. (magenta: PI; green: TUNEL+). BP, binucleate pollen; MMsp, mother microspore; T, tapetum; Tds, tetrads; PG, pollen grain; UMsp, uninucleate microspore. Scale bars are 100 μm (i), 2 cm and 500 μm (l left and right panel respectively), 2 cm (o), 50 μm (q–s). “N” indicates biological replicate (f), individual plant (j, m, n, p); data: mean ± SD (f, j) and box plots: median (central line); 25th and 75th percentiles (bounds); minima and maxima (whiskers) (m, n, p); P-values: Mann-Whitney U test (j), one-way ANOVA (m, n), Kruskal-Wallis test (p). Schematics in a and h created in BioRender; Chen, S. https://biorender.com/uc0y98j and https://biorender.com/8duox87 (2026).

Source data

Extended Data Fig. 3 Molecular features of SlRGFR6 and phenotypes of Slrgfr6 null mutants.

a, Phylogeny of RGFR families in tomato, rice and Arabidopsis. b, Heatmap depicting expression of seven SlRGFRs across various tomato tissues under normal conditions. EVM/MVM/LVM/TM/FM/SIM/SYM indicate the early vegetative/middle vegetative/late vegetative/transitional/floral/sympodial inflorescence/sympodial shoot meristem, respectively. c, Time-course expression profiles of SlRGFRs in tomato anthers under chilling. d, Anther section of SlRGFR6pro: GFP transgenic lines. e, CRISPR-Cas9 target design and allelic information of Slrgfr6 mutants. f–h, Representative images (f) and quantification of flowering time (g), and flower number per inflorescence (h) of wild-type and Slrgfr6 mutants grown at normal temperature. White asterisks: primary leaves; red arrowheads: primary inflorescences. i, Histology (top) and TUNEL (middle/bottom) of Slrgfr6 pollen development and tapetum PCD at normal temperature and after chilling (magenta: PI; green: TUNEL+). BP, binucleate pollen; DBP, degenerated binucleate pollen; DPG, degenerated pollen grain; MMC, microspore mother cell; T, tapetum; Tds, tetrads; PG, pollen grain; UMsp, uninucleate microspore. j–l, Effects of exogenous non-sulfated SlRGF applications on the fruit-setting rate (j, k) and pollen viability (j, l) of Slrgf9 Slrgf10 mutants under chilling conditions. m, Schematics of SlRGFR6 protein secondary structure. n, Subcellular localization of SlRGFR6 on plasma membranes of tobacco and tomato cells. GFP alone is used as a control. o, Expression and purification of SlRGFR6ECD. Red arrow: target protein. p, ITC detection of the binding of PSKs to SlRGFR6ECD. DP, differential power; ΔH, change in enthalpy. q–r, Molecular docking of SlRGFR6ECD with the SlRGF10. PTM: 0.88; IPTM: 0.96; pLDTT: 80.55. Scale bars are 50 μm (d), 2 cm and 50 μm (f left and right panel, respectively), 2 cm and 50 μm (j top and bottom panel, respectively), 50 μm (i), 10 μm (n). “N”: individual plant (g, h, k, l); data: box plots: median (central line); 25th and 75th percentiles (bounds); minima and maxima (whiskers) (g, h, k, l); P-values: Mann-Whitney U test (g, h), Kruskal-Wallis test (k, l).

Source data

Extended Data Fig. 4 Molecular features of SlSERKs and phenotypes of Slserk3a/3b null mutants.

a, Expression atlas of SlSERK family members in various tomato tissues. EVM/MVM/LVM/TM/FM/SIM/SYM indicate the early vegetative/middle vegetative/late vegetative/transitional/floral/sympodial inflorescence/sympodial shoot meristem, respectively. b, Co-IP showing SlRGF9/10 do not trigger SlRGFR6 homodimerization (PSK: negative control). c–e, BiFC assay showed SlRGF10-triggered interactions between SlRGFR6 and SlSERK1/3 A/3B. Representative images (c), quantification of fluorescence intensity (d) and western blot detection (e) of target proteins were shown. Non-sulfated SlRGF10 and scrambled peptides (SP) as negative controls. f, CRISPR-Cas9 target design for SlSERK3A/3B and allelic information of Slserk3a/3b double mutants. g, Phenotypes of wild-type and Slserk3a/3b double mutants under normal temperature. White arrows: fruit trusses/inflorescences. h–i, Alexander staining of pollen viability (h) and quantification (i) in wild-type, Slserk3a/3b homo-hetero double mutant grown at normal and chilling conditions. Yellow arrowheads: nonviable pollens. +/−, heterozygous mutant; −/−, homozygous mutant. j–k, Volcano plot and gene list of calcium signal-transduction genes in tomato anthers before /after chilling treatment. Two-sided DESeq2 with multiple-test correction (BH). Adjusted P < 0.05. Scale bars are 50 μm (c), 10 cm (g) and 50 μm (h). “N”: tobacco leaves (d) and individual plant (i); data: box plots: median (central line); 25th and 75th percentiles (bounds); minima and maxima (whiskers) (d, i); P-values: Mann-Whitney U test (d), Kruskal-Wallis test (i).

Source data

Extended Data Fig. 5 Expression pattern of SlCNGC genes and phenotypes of Slcngc7/8/16/18 mutants in tomato.

a, Expression atlas of SlCNGCs in tomato tissues. EVM/MVM/LVM/TM/FM/SIM/SYM indicate the early vegetative/middle vegetative/late vegetative/transitional/floral/sympodial inflorescence/sympodial shoot meristem, respectively. b, Time-course expression of SlCNGC7/8/16/18 in tomato anthers under chilling treatment. c, In situ mRNA hybridization showing the spatial expression pattern of SlCNGC7/8/16/18 in tomato anthers. Microspores and tapetum are indicated by red and yellow arrowheads, respectively. d, CRISPR-Cas9 gRNAs design for targeting SlCNGC7/8/16/18 and allelic information for Slcngc7/8/16/18 mutants. e–f, Representative images (e) and quantification (f) of fruit setting rates in Slcngc7/8 heterozygous mutants under normal and chilling conditions. Red arrowheads indicate flowers at tetrads and early uninucleate stages subject to chilling treatment that successfully developed into fruits. g–h, Slcngc16/18 homozygous mutants displayed defects in inflorescence architecture and flower development. Red arrowheads: primary inflorescences. −/−, homozygous mutant. Blots were processed in parallel. Scale bars are 50 μm (c), 2 cm (e), 10 cm (g), 1 cm (h). “N”: biological replicate (b); individual plant (f); data: mean ± SD (b) and box plots: median (central line); 25th and 75th percentiles (bounds); minima and maxima (whiskers) (f); P-values: two-tailed student’s t-test (b), Kruskal-Wallis test (f).

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Extended Data Fig. 6 Pollen development of Slcngc16/18 mutants.

a–b, Representative images (a) and quantification data (b) showing fruit setting rates in Slcngc16/18 heterozygous mutants under normal and chilling treatment. Red and white arrowheads indicate flowers at tetrads and early uninucleate stages subject to chilling treatment that successfully developed into fruits and aborted flowers, respectively. c–d, Histological analysis (c) and TUNEL assay (d) showing pollen development and tapetum PCD of Slcngc16/18 mutants under normal and chilling conditions (magenta: PI; green: TUNEL+). BP, binucleate pollen; DBP, degenerated binucleate pollen; DPG, degenerated pollen grain; MMC, microspore mother cell; T, tapetum; Tds, tetrads; PG, pollen grain; UMsp, uninucleate microspore. +/−, heterozygous mutant. e–g, Effects of exogenous SlRGF10 applications on fruit setting rates (e, f) and pollen viability (e, g) in wild-type and Slcngc16/18 heterozygous mutants under chilling. Scale bars are 2 cm (a), 50 μm (c–d), 2 cm, 50 μm respectively (e top and bottom panel, respectively). “N”: individual plant (b, f, g); data: box plots: median (central line); 25th and 75th percentiles (bounds); minima and maxima (whiskers) (b, f, g); P-values: Kruskal-Wallis test (b, f, g).

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Extended Data Fig. 7 In vitro phosphorylation of SlRGFR6 and SlCNGC16/18 and their plasma membrane–localization in HEK293T cells.

a, Co-IP showing SlRGFR6 interaction with SlCNGC16/18. SlERECTA: negative plasma membrane (PM)-localized control. b, Molecular docking of SlRGFR6 C-terminus (intracellular kinase) and adenosine triphosphate (ATP). PTM: 0.82; IPTM: 0.94. c, The Predicted Aligned Error (PAE) for AlphaFold prediction of SlRGFR6 C-terminus and ATP. pLDTT: 92.28. d, In vitro phosphorylation assays of purified C-terminal region of SlRGFR6 and SlRGFR6K797E mutant (kinase-dead). Autoradiography (left) and Coomassie brilliant blue staining (right) are shown. e, The SlRGFR6K797E fails to phosphorylate the N and C terminus of SlCNGC16/18. GST/MBP: negative controls. Blots were processed in parallel. f–h, PM-localization of SlRGFR6, SlSERK3A, SlCNGC16 and SlCNGC18 in HEK293T cells. Confocal images (f), colocalization analysis (g) and western blot detection of fusion proteins (h) were provided. The Manders’ Colocalization Coefficients (MMC) of the GFP and mCherry signal along the indicated dashed arrows were shown. Scale bars are 10 μm (f).

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Extended Data Fig. 8 Phenotypes of 35Spro: SlRGF9/10 T2 transgenic lines under normal conditions.

a–f, Phenotypes of wild-type and 35Spro: SlRGF9/10 T2 homozygous tomato plants under normal temperature at vegetative and flowering stages (a), along with quantification of main root length (b), number of 3 cm lateral roots (c), above-ground leaf area (d), leaves to primary inflorescence (e), and flower number per inflorescence (f). DAG, days after germination. g–m, Harvest phenotypes of wild-type and 35Spro: SlRGF9/10 transgenic tomato plants under normal temperature (g). Fruit truss (top), total fruits of first three inflorescences per plants (middle) and pollen viability (bottom). Yellow arrowheads: nonviable pollens. Along with quantification of SlRGF gene expression (h–i), fruit setting rate (j), individual fruit fresh weight (k), total fruit yield per plant (l) and pollen viability (m) for wild-type and SlRGF9/10 over-expression lines. Under normal temperature, the expression of SlRGF is almost undetectable in wild-type pollen, while it is significantly upregulated in over-expressing transgenic lines. Scale bars are 2 cm in (a), 2 cm, 2 cm, 50 μm respectively in (g top, middle and bottom panel, respectively). “N”: individual plant (b–f, j, l–m), biological replicate (h–i), fruit (k); data: mean ± SD (h–i) and box plots: median (central line); 25th and 75th percentiles (bounds); minima and maxima (whiskers) (b–f, j–m); P-values: two-tailed student’s t-test (h–i), one-way ANOVA (b–f, k–l), and Kruskal-Wallis test (j, m).

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Extended Data Fig. 9 Moderate up-regulation of SlRGF9/10 saves fruit-yield loss caused by chilling stress in solar greenhouse.

a, Daily daytime and nighttime temperature medians over a 50-d period of flower development under chilling conditions in winter greenhouse related to Fig. 5a–e. The gray dashed line indicates 12 °C. b, Daily daytime and nighttime temperature medians during flower development in the growing season under chilling conditions in early-spring greenhouse related to Extended Data Fig. 9c–n. The gray dashed lines indicate 12 °C and 20 °C. c–d, Representative images showing tomato populations (c), and fruit trusses of the first inflorescences (d) of wild-type and 35Spro: SlRGF9/10 T2 homozygous lines in the solar greenhouse under chilling stress in spring growth season. 1st, 2nd and 3rd indicate the inflorescence order. e–n, Quantification of fruit setting rate (e–g), individual fruit fresh weight (h), total fruit yield per inflorescence (i–k) and average plot yield (l–n) of first, second, and third inflorescences for wild-type and 35Spro: SlRGF9/10 transgenic lines. o, Daily daytime and nighttime temperature medians during flower development of the growing season facing chilling in the open field related to Supplementary Fig. 2a–e. The gray dashed line indicates 17 °C. Scale bars are 2 cm (d). “N”: individual plant and fruit (h), inflorescence (e–g, i–k), transplanting plot (l–n); data: box plots: median (central line); 25th and 75th percentiles (bounds); minima and maxima (whiskers) (e–n); P-values: Kruskal-Wallis test (e–g) and one-way ANOVA (h–n).

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Extended Data Fig. 10 Detection of sulfated SlRGF peptides in tomato anthers.

a–l, LC-HRMS detection of SlRGF peptide content within the anthers of wild-type and 35Spro: SlRGF9/10 transgenic tomato lines. Extracted ion chromatogram, isotopic distribution pattern and content quantification of SlRGF9 (a–c), non-sulfated SlRGF9 (d–f), SlRGF10 (g–i), and non-sulfated SlRGF10 (j–l) of the wild-type and 35Spro: SlRGF9/10 transgenic lines were shown. RT, retention time. Synthetic peptides as standards. m–n, Alexander staining (m) and quantification (n) showing pollen viability of wild-type and 35Spro: SlRGF9/10 transgenic lines under chilling stress. o–p, Confocal microscopy images (o) and quantification (p) of GCaMP6m fluorescence intensity showing Ca2+ concentration in living anthers of wild-type and SlRGF9/10 over-expression lines under chilling treatment. T, tapetum; UMsp, uninucleate microspore. Scale bars, 50 μm (m, o); “N”: technique replicate (c, f, i, l), individual plant (n), independent flower (p); data: box plots: median (central line); 25th and 75th percentiles (bounds); minima and maxima (whiskers) (n, p) and mean ± SD (c, f, i, l); P-values: Kruskal-Wallis test (n, p).

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Extended Data Fig. 11 Conserved OsRGF functions and phenotypes of Osrgf mutants and OsRGF10pro: OsRGF10 transgenic lines in rice.

a, Phylogeny of RGF families in tomato, rice, wheat (Triticum aestivum), maize (Zea mays), soybean (Glycine max), sorghum (Sorghum bicolor) and millet (Setaria italica). b, CRISPR-Cas9 gRNAs design for targeting OsRGF8/10 and allelic information for Osrgf8/10 mutants. c–d, Floret (left) and pollen (right) morphology (c) and quantification of pollen viability (d) of rice c.v. ZH11 and Osrgf mutants under chilling. e, Quantification of OsRGF10 expression in OsRGF10pro: OsRGF10 transgenic lines under 15 °C chilling. f, Daily daytime and nighttime temperature medians before the day of heading related to Extended Data Fig. 11g–r. g–j, Representative images (g) and quantification of heading stage (h), plant height (i) and tiller number (j) of ZH11 and OsRGF10pro: OsRGF10 transgenic lines grown at normal temperature in northern China (39.9°N, 116.3°E). k–p, Representative images of panicles and spikelets (k) and quantification of primary spike branches (l), grains per panicle (m), seed setting rate (n), 1,000-grain weight (o), and grain yield per plant (p) of ZH11 and OsRGF10pro: OsRGF10 transgenic lines grown. White asterisks: panicle primary branches. q–r, Florets and pollen morphology (q) and quantification (r) for ZH11 and OsRGF10pro: OsRGF10 transgenic lines. Scale bars are 1 mm, 1 mm, 1 mm and 100 μm respectively (c from left to right). 10 cm (g), 3 cm (k), 1 mm, 1 mm, 100 μm respectively (q top, middle and bottom panel). “N”: biological replicate (e), individual plant (d, h–j, o–p, r), panicle (l–n), data: mean ± SD (e); relative percentage (h), box plots: median (central line); 25th and 75th percentiles (bounds); minima and maxima (whiskers) (d, i–j, l–p, r). P-values: Kruskal-Wallis tests (d, n, r), two-tailed student’s t-test (e), one-way ANOVA (i–j, l–m, o–p).

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Extended Data Fig. 12 Plot yield trials of OsRGF10pro: OsRGF10 T2 transgenic rice in paddy field under chilling in northern and southern China.

a, Daily daytime and nighttime temperature medians before heading related to Extended Data Fig. 12a–j. b, Performance of ZH11 and OsRGF10pro: OsRGF10 T2 homozygous plants at late grain-filling stage under chilling stress in paddy fields in northern China (39.9°N, 116.3°E). c–e, Representative images of thirty-five harvested plants (c), along with quantification of heading stage (d), tiller number (e), seed setting rate (f), 1,000-grain weight (g), grain yield per plant (h), grain yield per plot (i) and pollen viability (j) of ZH11 and OsRGF10pro: OsRGF10 transgenic lines under chilling. k, Comparison of full and empty grain per panicle among ZH11 and OsRGF10pro: OsRGF10 transgenic plants under chilling stress in in southern China (22.0°N, 100.5°E). White asterisk: panicle primary branch. l, Daily daytime and nighttime temperature medians under chilling conditions before heading related to Extended Data Fig. 12k–s. m–s, Quantification of tiller number (m), panicle primary branches (n), grains per panicle (o), seed setting rate (p), 1,000-grain weight (q), grain yield per plant (r), and grain yield per plot (s) of ZH11 and OsRGF10pro: OsRGF10 transgenic lines under chilling stress. Scale bars are 2 cm (c, i). “N”: individual plant (d–h, j, m–r), transplanting plot (i, s), data: relative percentage (d), box plots: median (central line); 25th and 75th percentiles (bounds); minima and maxima (whiskers) (e–j, m–r) and mean ± SD (s); P-values: one-way ANOVA (e, g–i, m–o, q–s), and Kruskal-Wallis test (f, j, p).

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Supplementary information

Supplementary Information (download PDF )

Supplementary Methods describe the plant growth conditions, chilling treatment and yield trials. Supplementary Figs. 1−3 contain peptide and LRR-RLK sequence alignments and the yield trial results. Supplementary Fig. 4 contains uncropped blot images.

Reporting Summary (download PDF )

Supplementary Tables 1–3 (download XLSX )

Supplementary Table 1 contains the SlSSPs annotated in this study. Supplementary Table 2 contains the SlSSPs that were undetectable at normal temperature but were chilling-induced in tomato anthers. Supplementary Table 3 contains the primers used in this study.

Peer Review file (download PDF )

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Chen, S., Zou, Y., Cui, H. et al. Cold-induced peptide signalling secures pollen resilience and crop yield. Nature (2026). https://doi.org/10.1038/s41586-026-10603-7

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