# Molecular mechanisms and breeding strategies for heat tolerance in vegetable crops under global warming

**Authors:** Yanlong Li, Xi Zhang, Chan Xia, Ting Wu, Yuyu Gao, Lingen Zeng, Zhuoxuan Wu, Xiongze Dai, Fang Yuan, Feng Liu, Sha Yang, Xuexiao Zou

PMC · DOI: 10.1093/hr/uhaf309 · Horticulture Research · 2025-11-06

## TL;DR

This paper reviews how extreme heat affects vegetable crops and proposes breeding strategies to develop heat-tolerant varieties for food security under climate change.

## Contribution

The paper introduces six strategic pathways for breeding heat-resilient vegetables based on molecular mechanisms and interdisciplinary approaches.

## Key findings

- Heat stress disrupts vegetable development from germination to reproduction, causing morphological and physiological impairments.
- Molecular mechanisms like HSFs, HSPs, and Ca2+-ROS signaling contribute to thermotolerance in vegetables.
- Strategies such as CRISPR editing, epigenetic memory, and synthetic promoters are proposed to enhance heat resilience.

## Abstract

Extreme heat driven by climate change poses a catastrophic threat to global vegetable production, undermining nutritional security because of the heightened physiological sensitivity and succulent tissues of these crops. This review synthesizes the multistage impacts of heat stress across critical developmental phases—from germination to reproduction—emphasizing morphological impairments (such as leaf wilting and floral abortion) and physiological disruptions (including photosynthetic inhibition and oxidative damage). We systematically dissect thermotolerance mechanisms in vegetables, highlighting transcriptional reprogramming by HSFs, WRKY, and NAC transcription factors; chaperone-mediated proteostasis via HSPs; epigenetic remodeling; Ca2+-ROS signaling pathways; and the role of phase separation dynamics. Importantly, we propose six strategic pathways to develop heat-resilient vegetables: harnessing natural variation through pan-genome-driven allele mining; employing biotechnological interventions such as CRISPR-mediated editing and synthetic promoters; engineering multistress tolerance by targeting conserved ‘core response’ pathways; exploiting epigenetic memory to achieve transgenerational resilience; optimizing source-sink dynamics with ‘’Climate-Responsive Carbon Optimization; and applying plant growth regulators and nanotechnology to enhance thermotolerance. Together, these strategies chart a clear roadmap for climate-smart vegetable breeding and call for interdisciplinary collaboration to translate molecular discoveries into practical breeding approaches for sustainable food systems under escalating thermal extremes.

## Linked entities

- **Genes:** WRKY (probable WRKY transcription factor 33) [NCBI Gene 103865671], XK (X-linked Kx blood group antigen, Kell and VPS13A binding protein) [NCBI Gene 7504], hsp70-1 (heat shock protein 70-1) [NCBI Gene 3879515], CA2 (carbonic anhydrase 2) [NCBI Gene 760]

## Full-text entities

- **Genes:** APX [NCBI Gene 778224], peroxidase [NCBI Gene 543959], APX1 [NCBI Gene 778223], CaM [NCBI Gene 101494641], RuvB-like 2 [NCBI Gene 101262595], FESOD (iron superoxide dismutase) [NCBI Gene 544259] {aka Fe-SODle, sodb}, RBOHD (respiratory burst oxidase homologue D) [NCBI Gene 834842] {aka ATRBOHD, MCA23.25, MCA23_25, RESPIRATORY BURST OXIDASE, respiratory burst oxidase homologue D}, SIZ1 [NCBI Gene 101252253], P5CS [NCBI Gene 544281], Coi1 (coronatine-insensitive 1) [NCBI Gene 543911], Multiprotein Bridging Factor 1c [NCBI Gene 101245564], MYB [NCBI Gene 544113], MiR164a [NCBI Gene 104794943], NUA (nuclear pore anchor) [NCBI Gene 844268] {aka AtTPR, TRANSLOCATED PROMOTER REGION, YUP8H12R.12, YUP8H12R_12, nuclear pore anchor}, HSFA2 (heat shock transcription factor A2) [NCBI Gene 817155] {aka ATHSFA2, T19L18.4, T19L18_4, heat shock transcription factor A2}, PIF4 (phytochrome interacting factor 4) [NCBI Gene 818903] {aka AtPIF4, MFL8.13, MFL8_13, SRL2, phytochrome interacting factor 4}, HSP70 [NCBI Gene 100134914], ABA1 (zeaxanthin epoxidase (ZEP) (ABA1)) [NCBI Gene 836838] {aka ABA DEFICIENT 1, ARABIDOPSIS THALIANA ABA DEFICIENT 1, ARABIDOPSIS THALIANA ZEAXANTHIN EPOXIDASE, ATABA1, ATZEP, IBS3}, ELF3 (hydroxyproline-rich glycoprotein family protein) [NCBI Gene 817134] {aka EARLY FLOWERING 3, F17H15.25, PYK20}, PSBO (oxygen-evolving enhancer protein 1, chloroplastic) [NCBI Gene 544299] {aka OEE1}, GID1b-1 (gibberellin receptor GID1b-1) [NCBI Gene 100736493] {aka GID1, SlGID1b-1}, Diacylglycerol Kinase [NCBI Gene 544171], SGT1 [NCBI Gene 101247681], PHYB2 (phytochrome B2) [NCBI Gene 101264523] {aka PHYB/D}, LOC101254720 (abscisic acid 8'-hydroxylase 3) [NCBI Gene 101254720] {aka SlCYP707A3}, actin [NCBI Gene 101260631], HSP90 [NCBI Gene 101260143], NCED6 (nine-cis-epoxycarotenoid dioxygenase 6) [NCBI Gene 111879595] {aka LsNCED4, NCED4}, SNCA (synuclein alpha) [NCBI Gene 6622] {aka NACP, PARK1, PARK4, PD1}, Mitogen-activated protein kinase [NCBI Gene 543792], WRKY [NCBI Gene 100191122], LIN5 [NCBI Gene 544241], glyceraldehyde-3-phosphate dehydrogenase [NCBI Gene 100736499], RBOH1 (NADPH oxidase) [NCBI Gene 543571] {aka gp91}, HSP70 (heat shock protein 70) [NCBI Gene 820438] {aka ARABIDOPSIS HEAT SHOCK PROTEIN 70, ATHSP70, heat shock protein 70}, MAPKK [NCBI Gene 543948], PLDa1 (phospholipase PLDa1) [NCBI Gene 543819], NAC1 (NAC domain protein) [NCBI Gene 543917] {aka NC1, SlNAC1}, SNF1-Related Protein Kinase [NCBI Gene 543870], MPK2 [NCBI Gene 543879], HSP101 (heat shock protein 101) [NCBI Gene 843771] {aka ATHSP101, F1O17.2, F1O17_2, HEAT SHOCK PROTEIN ATHSP101, HOT1, heat shock protein 101}, CURL3 (brassinosteroid LRR receptor kinase) [NCBI Gene 101261320] {aka BRI1, SR160, SlBRI1, tBRI1}, catalase [NCBI Gene 101513499], calmodulin [NCBI Gene 101489805], HsfA1a (heat stress transcription factor HsfA1a) [NCBI Gene 101263626] {aka hsf8}, psbA [NCBI Gene 9073046], TPL (Transducin family protein / WD-40 repeat family protein) [NCBI Gene 838144] {aka F7H2.9, F7H2_9, TOPLESS, WSIP1, WUS-INTERACTING PROTEIN 1}, StMPK1 [NCBI Gene 102599364], PAL5 (phenylalanine ammonia-lyase 5) [NCBI Gene 101244220] {aka PAL}
- **Diseases:** infectious (MESH:D003141), floral abnormalities (MESH:D000014), drought (MESH:C536747), cardiovascular disorders (MESH:D002318), PrLD (MESH:D017096), respiratory dysfunction (MESH:D012131), HS (MESH:D018882), black heart (MESH:D007898), reproductive (MESH:D060737), diabetes (MESH:D003920), water deficits (MESH:D000069578), PSII impairment (MESH:D060825), reproductive failure (MESH:D051437)
- **Chemicals:** sucrose (MESH:D013395), lipid (MESH:D008055), CO2 (MESH:D002245), Cytokinin (MESH:D003583), polyamine (MESH:D011073), ATP (MESH:D000255), Auxin (MESH:D007210), ROS (MESH:D017382), calcium (MESH:D002118), JA (MESH:C011006), 5-azacytidine (MESH:D001374), flavonoids (MESH:D005419), membrane lipid (MESH:D008563), anthocyanin (MESH:D000872), GA (MESH:D005708), MT (MESH:D008550), H2O2 (MESH:D006861), O2- (MESH:D013481), Aux (-), fatty acid (MESH:D005227), carbohydrate (MESH:D002241), starch (MESH:D013213), amino acid (MESH:D000596), Gibberellin (MESH:D005875), carotenoids (MESH:D002338), CeO2 (MESH:C030583), lycopene (MESH:D000077276), Water (MESH:D014867), hydroxyl radicals (MESH:D017665), BR (MESH:D060406), SA (MESH:D020156), sugar (MESH:D000073893), singlet oxygen (MESH:D026082), ABA (MESH:D000040), Pr (MESH:D011221), Carbon (MESH:D002244), phosphatidic acid (MESH:D010712), chlorophyll (MESH:D002734), nitrogen (MESH:D009584), Ethylene (MESH:C036216), polysaccharides (MESH:D011134)
- **Species:** Cucumis sativus (cucumber, species) [taxon 3659], Oryza sativa (Asian cultivated rice, species) [taxon 4530], Brassica oleracea var. botrytis (cauliflower, varietas) [taxon 3715], Brassica oleracea var. italica (asparagus broccoli, varietas) [taxon 36774], Brassica rapa subsp. pekinensis (bai cai, subspecies) [taxon 51351], Asparagus officinalis (garden asparagus, species) [taxon 4686], Solanum tuberosum (potatoes, species) [taxon 4113], Hemerocallis citrina (species) [taxon 1249515], Hemerocallis hybrid cultivar (daylily, species) [taxon 29711], Triticum aestivum (bread wheat, species) [taxon 4565], Lathyrus oleraceus (garden pea, species) [taxon 3888], Allium cepa (onion, species) [taxon 4679], Daucus carota (carrot, species) [taxon 4039], Capsicum annuum (sweet pepper, species) [taxon 4072], Phaseolus vulgaris (common bean, species) [taxon 3885], Arabidopsis thaliana (mouse-ear cress, species) [taxon 3702], Solanum melongena (aubergine, species) [taxon 4111], Solanum lycopersicum (tomato, species) [taxon 4081], Powellomyces sp. EA (species) [taxon 252690], Homo sapiens (human, species) [taxon 9606], Zea mays (maize, species) [taxon 4577], Capsicum annuum var. annuum (jalapeno pepper, varietas) [taxon 40321], Apium graveolens Dulce Group (celery, no rank) [taxon 117781], Brassica oleracea var. capitata (cabbage, varietas) [taxon 3716], Brassica oleracea (wild cabbage, species) [taxon 3712], Abelmoschus esculentus (lady's fingers, species) [taxon 455045], Raphanus sativus (radish, species) [taxon 3726], Setaria viridis (species) [taxon 4556], Arachis hypogaea (goober, species) [taxon 3818], Spinacia oleracea (spinach, species) [taxon 3562], Cicer arietinum (chickpea, species) [taxon 3827], Glycine max (soybean, species) [taxon 3847], Apium graveolens (species) [taxon 4045], Capsicum frutescens (bird pepper, species) [taxon 4073], Lactuca sativa (cultivated lettuce, species) [taxon 4236], Gossypium hirsutum (American cotton, species) [taxon 3635]
- **Mutations:** C-45 C, C-20 C, C-28 C, C-25 C, G/A

## Full text

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## Figures

4 figures with captions in the complete paper: https://tomesphere.com/paper/PMC12936445/full.md

## References

306 references — full list in the complete paper: https://tomesphere.com/paper/PMC12936445/full.md

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Source: https://tomesphere.com/paper/PMC12936445