{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T05:01:06Z","timestamp":1790226066109,"version":"4.1.0"},"reference-count":46,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T00:00:00Z","timestamp":1790208000000},"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:sec>\n                    <jats:title>Introduction<\/jats:title>\n                    <jats:p>\n                      Understanding the molecular mechanisms underlying fruit color variation is crucial for citrus breeding. The variegated \u2018Gonggan\u2019 mandarin (\n                      <jats:italic>Citrus reticulata<\/jats:italic>\n                      Blanco) exhibits distinct phenotypic differences compared to the normal cultivar. However, the physiological and genetic basis for this variation remains unclear.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>In this study, we integrated physiological measurements, liquid chromatography-mass spectrometry (LC-MS), and transcriptome sequencing to characterize the pericarp of the variegated \u2018Gonggan\u2019 mandarin during early fruit development stage.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      Compared with the normal \u2018Gonggan\u2019 mandarin, lightness and chromaticity values of the pericarp color exhibited significantly elevated in the variegated \u2018Gonggan\u2019 mandarin. Physiological assays revealed a drastic reduction in chlorophyll a and b contents (by 75.76% and 58.33%, respectively) and a moderate increase in total carotenoids (28.61%) in the variegated \u2018Gonggan\u2019 mandarin. Notably, phytoene accumulation in the pericarp of the variegated \u2018Gonggan\u2019 mandarin was 44.35-fold higher than that in the normal fruit, while metabolites in the middle and downstream pathway of carotenoid biosynthesis such as \u03b2-carotene and lutein were significantly reduced. Transcriptome analysis identified 1,438 differentially expressed genes (DEGs), which were predominantly enriched in metabolic pathways, secondary metabolite biosynthesis and photosynthesis. Specifically, 3 chlorophyll degradation genes encoding chlorophyll b reductase gene, chlorophyllase and SGR protein, 23 photosynthesis genes (\n                      <jats:italic>PsaD<\/jats:italic>\n                      ,\n                      <jats:italic>PsaE<\/jats:italic>\n                      ,\n                      <jats:italic>PsbO<\/jats:italic>\n                      ,\n                      <jats:italic>etc<\/jats:italic>\n                      .), and 6 photosynthesis-antenna proteins (\n                      <jats:italic>Lhca1<\/jats:italic>\n                      ,\n                      <jats:italic>Lhca2<\/jats:italic>\n                      ,\n                      <jats:italic>Lhcb1<\/jats:italic>\n                      ,\n                      <jats:italic>etc<\/jats:italic>\n                      .) were significantly upregulated. Furthermore, 13 DEGs associated with plant hormone signal transduction exhibited differential expression patterns.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Discussion<\/jats:title>\n                    <jats:p>Our findings demonstrate that the disruption of the chlorophyll degradation pathway and carotenoid biosynthesis pathway are the primary causes of the pericarp color variation in the variegated \u2018Gonggan\u2019 mandarin. This study provides novel insights into the regulatory network governing pericarp coloration and establishes a foundation for future molecular breeding strategies targeting citrus appearance quality.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fpls.2026.1960602","type":"journal-article","created":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T04:30:35Z","timestamp":1790224235000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Integrated metabolomic and transcriptomic analysis of the mechanism underlying pericarp variegation in Citrus reticulata \u2018Gonggan\u2019"],"prefix":"10.3389","volume":"17","author":[{"given":"Congyi","family":"Zhu","sequence":"first","affiliation":[{"name":"Institute of Fruit Tree Research, Guangdong Academy of Agricultural Sciences, Key Laboratory of South Subtropical Fruit Biology and Genetic Resource Utilization (MOA), Guangdong Province Key Laboratory of Science and Technology Research on Fruit Tree","place":["Guangzhou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pingzhi","family":"Wu","sequence":"additional","affiliation":[{"name":"Institute of Fruit Tree Research, Guangdong Academy of Agricultural Sciences, Key Laboratory of South Subtropical Fruit Biology and Genetic Resource Utilization (MOA), Guangdong Province Key Laboratory of Science and Technology Research on Fruit Tree","place":["Guangzhou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yibo","family":"Hu","sequence":"additional","affiliation":[{"name":"School of Life Sciences, Zhaoqing University","place":["Zhaoqing, China"]},{"name":"Deqing Citrus Research Institute","place":["Zhaoqing, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ruimin","family":"Zhang","sequence":"additional","affiliation":[{"name":"Institute of Fruit Tree Research, Guangdong Academy of Agricultural Sciences, Key Laboratory of South Subtropical Fruit Biology and Genetic Resource Utilization (MOA), Guangdong Province Key Laboratory of Science and Technology Research on Fruit Tree","place":["Guangzhou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hongqi","family":"Xia","sequence":"additional","affiliation":[{"name":"Institute of Fruit Tree Research, Guangdong Academy of Agricultural Sciences, Key Laboratory of South Subtropical Fruit Biology and Genetic Resource Utilization (MOA), Guangdong Province Key Laboratory of Science and Technology Research on Fruit Tree","place":["Guangzhou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhengyan","family":"Fan","sequence":"additional","affiliation":[{"name":"Institute of Fruit Tree Research, Guangdong Academy of Agricultural Sciences, Key Laboratory of South Subtropical Fruit Biology and Genetic Resource Utilization (MOA), Guangdong Province Key Laboratory of Science and Technology Research on Fruit Tree","place":["Guangzhou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pengtao","family":"Yue","sequence":"additional","affiliation":[{"name":"Institute of Fruit Tree Research, Guangdong Academy of Agricultural Sciences, Key Laboratory of South Subtropical Fruit Biology and Genetic Resource Utilization (MOA), Guangdong Province Key Laboratory of Science and Technology Research on Fruit Tree","place":["Guangzhou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chuanhong","family":"Ren","sequence":"additional","affiliation":[{"name":"Institute of Fruit Tree Research, Guangdong Academy of Agricultural Sciences, Key Laboratory of South Subtropical Fruit Biology and Genetic Resource Utilization (MOA), Guangdong Province Key Laboratory of Science and Technology Research on Fruit Tree","place":["Guangzhou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peihua","family":"Lin","sequence":"additional","affiliation":[{"name":"Deqing Citrus Research Institute","place":["Zhaoqing, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sai","family":"Xu","sequence":"additional","affiliation":[{"name":"Institute of Facility Agriculture, Guangdong Academy of Agricultural Sciences","place":["Guangzhou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yongjing","family":"Huang","sequence":"additional","affiliation":[{"name":"Institute of Fruit Tree Research, Guangdong Academy of Agricultural Sciences, Key Laboratory of South Subtropical Fruit Biology and Genetic Resource Utilization (MOA), Guangdong Province Key Laboratory of Science and Technology Research on Fruit Tree","place":["Guangzhou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,9,24]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1104\/pp.24.1.1","article-title":"Copper enzymes in isolated chloroplasts. 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