{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,2]],"date-time":"2026-09-02T11:13:02Z","timestamp":1788347582396,"version":"build-2803163510"},"reference-count":24,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,5,21]],"date-time":"2026-05-21T00:00:00Z","timestamp":1779321600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Plant Sci."],"abstract":"<jats:p>\n                    As an economically important vegetable species, bitter gourd\n                    <jats:italic>(Momordica charantia<\/jats:italic>\n                    L.) is widely cultivated around the globe. However, its production is frequently threatened by late spring coldness, which severely impairs seedling establishment, delays flowering and fruit set, and reduces both yield and quality. Unraveling mechanisms of cold resistance in bitter gourd is therefore critical for breeding resilient cultivars and expanding its planting range. Here, we found a cold-inducible\n                    <jats:italic>CIPK<\/jats:italic>\n                    family gene,\n                    <jats:italic>McCIPK6<\/jats:italic>\n                    , from bitter gourd via transcriptome screening, and systematically characterized its function in cold stress responses using phenotypic, physiological, and molecular approaches. Expression analysis revealed that\n                    <jats:italic>McCIPK6<\/jats:italic>\n                    was rapidly and strongly induced in the cold-tolerant genotype \u20180208\u2019 (R) under low temperature, whereas the cold-sensitive line \u20182206\u2019 (S) showed a delayed and dysregulated expression pattern. Mitochondrial localization analysis revealed that McCIPK6 is a mitochondria\u2212targeted CIPK kinase. Heterologous overexpression of\n                    <jats:italic>McCIPK6<\/jats:italic>\n                    in Arabidopsis significantly enhanced cold tolerance by simultaneously promoting mitochondrial ROS scavenging and activating the\n                    <jats:italic>CBF<\/jats:italic>\n                    \u2010dependent cold signaling pathway. These effects were reflected by improved survival rates, reduced MDA accumulation, lower ROS levels, higher antioxidant enzyme activities (SOD, POD, CAT), and upregulated expression of\n                    <jats:italic>CBF<\/jats:italic>\n                    regulation genes as well as antioxidant genes. Further verification via yeast two-hybrid and luciferase complementation experiments revealed that McCIPK6 physically interacts with McCBL1, a core calcium sensor in bitter gourd. Collectively, our data establish that\n                    <jats:italic>McCIPK6<\/jats:italic>\n                    functions as a positive regulator of cold tolerance by regulating ROS scavenging and the\n                    <jats:italic>CBF<\/jats:italic>\n                    -dependent cold signaling cascade, likely through forming a functional complex with McCBL1. This study reveals a novel regulatory module underlying cold adaptation in bitter gourd and offers a valuable candidate gene for the genetic enhancement of cold resistance in cucurbit crops.\n                  <\/jats:p>","DOI":"10.3389\/fpls.2026.1856408","type":"journal-article","created":{"date-parts":[[2026,5,21]],"date-time":"2026-05-21T04:52:52Z","timestamp":1779339172000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Characterization of McCIPK6, a mitochondrial protein, positively regulating cold stress tolerance in bitter gourd (Momordica charantia L.)"],"prefix":"10.3389","volume":"17","author":[{"given":"Xuetong","family":"Yang","sequence":"first","affiliation":[{"name":"Institute of Vegetables and Flowers, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]},{"name":"Jiangxi Key Laboratory of Horticultural Crops (Fruit, Vegetable & Tea) Breeding, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]},{"name":"Jiangxi Engineering Research Center of Vegetable Molecular Breeding, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kai","family":"Wang","sequence":"additional","affiliation":[{"name":"Institute of Vegetables and Flowers, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]},{"name":"Jiangxi Key Laboratory of Horticultural Crops (Fruit, Vegetable & Tea) Breeding, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]},{"name":"Jiangxi Engineering Research Center of Vegetable Molecular Breeding, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuanyuan","family":"Xie","sequence":"additional","affiliation":[{"name":"Institute of Vegetables and Flowers, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]},{"name":"Jiangxi Key Laboratory of Horticultural Crops (Fruit, Vegetable & Tea) Breeding, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]},{"name":"Jiangxi Engineering Research Center of Vegetable Molecular Breeding, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Feng","family":"Guan","sequence":"additional","affiliation":[{"name":"Institute of Vegetables and Flowers, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]},{"name":"Jiangxi Key Laboratory of Horticultural Crops (Fruit, Vegetable & Tea) Breeding, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]},{"name":"Jiangxi Engineering Research Center of Vegetable Molecular Breeding, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bo","family":"Shi","sequence":"additional","affiliation":[{"name":"Institute of Vegetables and Flowers, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]},{"name":"Jiangxi Key Laboratory of Horticultural Crops (Fruit, Vegetable & Tea) Breeding, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]},{"name":"Jiangxi Engineering Research Center of Vegetable Molecular Breeding, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinjian","family":"Wan","sequence":"additional","affiliation":[{"name":"Institute of Vegetables and Flowers, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]},{"name":"Jiangxi Key Laboratory of Horticultural Crops (Fruit, Vegetable & Tea) Breeding, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]},{"name":"Jiangxi Engineering Research Center of Vegetable Molecular Breeding, Jiangxi Academy of Agricultural Sciences","place":["Nanchang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,5,21]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.3390\/genes10060446","article-title":"Ectopic expression of cold responsive LlaCIPK gene enhances cold stress tolerance in Nicotiana tabacum","volume":"10","author":"Aslam","year":"2019","journal-title":"Genes (Basel)"},{"key":"B2","doi-asserted-by":"publisher","DOI":"10.1016\/j.plantsci.2022.111284","article-title":"AcCIPK5, a pineapple CBL-interacting protein kinase, confers salt, osmotic and cold stress tolerance in transgenic Arabidopsis","volume":"320","author":"Aslam","year":"2022","journal-title":"Plant Sci."},{"key":"B3","doi-asserted-by":"publisher","first-page":"6211","DOI":"10.1093\/jxb\/ers273","article-title":"The Brassica napus calcineurin B-Like 1\/CBL-interacting protein kinase 6 (CBL1\/CIPK6) component is involved in the plant response to abiotic stress and ABA signalling","volume":"63","author":"Chen","year":"2012","journal-title":"J. Exp. Bot."},{"key":"B4","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1007\/s11103-024-01417-0","article-title":"The role of CBL-CIPK signaling in plant responses to biotic and abiotic stresses","volume":"114","author":"Chen","year":"2024","journal-title":"Plant Mol. Biol."},{"key":"B5","doi-asserted-by":"publisher","first-page":"4759","DOI":"10.1007\/s11033-013-2572-9","article-title":"Arabidopsis CBL-interacting protein kinase (CIPK6) is involved in plant response to salt\/osmotic stress and ABA","volume":"40","author":"Chen","year":"2013","journal-title":"Mol. Biol. Rep."},{"key":"B6","doi-asserted-by":"publisher","first-page":"2748","DOI":"10.1105\/tpc.113.113530","article-title":"The tomato calcium sensor Cbl10 and its interacting protein kinase Cipk6 define a signaling pathway in plant immunity","volume":"25","author":"de la Torre","year":"2013","journal-title":"Plant Cell"},{"key":"B7","doi-asserted-by":"publisher","first-page":"236","DOI":"10.1104\/pp.19.01368","article-title":"The calcium sensor CBL2 and its interacting kinase CIPK6 are involved in plant sugar homeostasis via interacting with tonoplast sugar transporter TST2","volume":"183","author":"Deng","year":"2020","journal-title":"Plant Physiol."},{"key":"B8","doi-asserted-by":"publisher","first-page":"367","DOI":"10.1111\/ppl.12046","article-title":"Ectopic expression of wheat TaCIPK14, encoding a calcineurin B-like protein-interacting protein kinase, confers salinity and cold tolerance in tobacco","volume":"149","author":"Deng","year":"2013","journal-title":"Physiol. Plant"},{"key":"B9","doi-asserted-by":"publisher","first-page":"4786","DOI":"10.1111\/pce.15070","article-title":"CIPK11 phosphorylates GSTU23 to promote cold tolerance in Camellia sinensis","volume":"47","author":"Di","year":"2024","journal-title":"Plant Cell Environ."},{"key":"B10","doi-asserted-by":"publisher","first-page":"1116","DOI":"10.1038\/cr.2011.50","article-title":"Calcium-dependent modulation and plasma membrane targeting of the AKT2 potassium channel by the CBL4\/CIPK6 calcium sensor\/protein kinase complex","volume":"21","author":"Held","year":"2011","journal-title":"Cell Res."},{"key":"B11","doi-asserted-by":"publisher","first-page":"2142","DOI":"10.1105\/tpc.105.032508","article-title":"Calcium: a central regulator of plant growth and development","volume":"17","author":"Hepler","year":"2005","journal-title":"Plant Cell"},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.3390\/ijms25095043","article-title":"Molecular mechanisms of CBL-CIPK signaling pathway in plant abiotic stress tolerance and hormone crosstalk","volume":"25","author":"Kaya","year":"2024","journal-title":"Int. J. Mol. Sci."},{"key":"B13","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1104\/pp.103.033068","article-title":"Calcium sensors and their interacting protein kinases: genomics of the Arabidopsis and rice CBL-CIPK signaling networks","volume":"134","author":"Kolukisaoglu","year":"2004","journal-title":"Plant Physiol."},{"key":"B14","doi-asserted-by":"publisher","first-page":"199","DOI":"10.1093\/abbs\/gmx140","article-title":"Ectopic expression of BdCIPK31 confers enhanced low-temperature tolerance in transgenic tobacco plants","volume":"50","author":"Luo","year":"2018","journal-title":"Acta Biochim. Biophys. Sin. (Shanghai)"},{"key":"B15","doi-asserted-by":"publisher","first-page":"1655","DOI":"10.1093\/jxb\/erab505","article-title":"The CBL-interacting protein kinase CaCIPK13 positively regulates defence mechanisms against cold stress in pepper","volume":"73","author":"Ma","year":"2022","journal-title":"J. Exp. Bot."},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.3390\/ijms21165668","article-title":"The CBL-CIPK pathway in plant response to stress signals","volume":"21","author":"Ma","year":"2020","journal-title":"Int. J. Mol. Sci."},{"key":"B17","doi-asserted-by":"publisher","DOI":"10.1155\/2015\/493191","article-title":"The calcium sensor CBL-CIPK is involved in plant's response to abiotic stresses","volume":"2015","author":"Manik","year":"2015","journal-title":"Int. J. Genomics"},{"key":"B18","doi-asserted-by":"publisher","first-page":"604","DOI":"10.1016\/j.tplants.2020.01.009","article-title":"The CBL-CIPK calcium signaling network: Unified paradigm from 20 years of discoveries","volume":"25","author":"Tang","year":"2020","journal-title":"Trends Plant Sci."},{"key":"B19","doi-asserted-by":"publisher","first-page":"539","DOI":"10.1007\/s00425-011-1522-9","article-title":"An ER-targeted calcium-binding peptide confers salt and drought tolerance mediated by CIPK6 in Arabidopsis","volume":"235","author":"Tsou","year":"2012","journal-title":"Planta"},{"key":"B20","doi-asserted-by":"publisher","first-page":"1416","DOI":"10.1104\/pp.107.101295","article-title":"Characterization of stress-responsive CIPK genes in rice for stress tolerance improvement","volume":"144","author":"Xiang","year":"2007","journal-title":"Plant Physiol."},{"key":"B21","doi-asserted-by":"publisher","first-page":"420","DOI":"10.1186\/s12870-022-03809-0","article-title":"Evolution, gene expression, and protein\u2013protein interaction analyses identify candidate CBL-CIPK signalling networks implicated in stress responses to cold and bacterial infection in citrus","volume":"22","author":"Xiao","year":"2022","journal-title":"BMC Plant Biol."},{"key":"B22","doi-asserted-by":"publisher","first-page":"3440","DOI":"10.3390\/plants14223440","article-title":"The bitter gourd transcription factor McNAC087 confers cold resistance in transgenic Arabidopsis","volume":"14","author":"Yang","year":"2025","journal-title":"Plants"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.3390\/ijms21010110","article-title":"Identification and functional analysis of tomato CIPK gene family","volume":"21","author":"Zhang","year":"2019","journal-title":"Int. J. Mol. Sci."},{"key":"B24","doi-asserted-by":"publisher","DOI":"10.1093\/hr\/uhae289","article-title":"NpCIPK6-NpSnRK1 module facilitates intersubgeneric hybridization barriers in water lily (Nymphaea) by reducing abscisic acid content","volume":"12","author":"Zhou","year":"2025","journal-title":"Hortic. Res."}],"updated-by":[{"DOI":"10.3389\/fpls.2026.1973395","type":"correction","label":"Correction","source":"publisher","updated":{"date-parts":[[2026,9,2]],"date-time":"2026-09-02T00:00:00Z","timestamp":1788307200000}}],"container-title":["Frontiers in Plant Science"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1856408\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,2]],"date-time":"2026-09-02T10:08:51Z","timestamp":1788343731000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1856408\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5,21]]},"references-count":24,"alternative-id":["10.3389\/fpls.2026.1856408"],"URL":"https:\/\/doi.org\/10.3389\/fpls.2026.1856408","relation":{},"ISSN":["1664-462X"],"issn-type":[{"value":"1664-462X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5,21]]},"article-number":"1856408"}}