{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,28]],"date-time":"2026-05-28T05:05:24Z","timestamp":1779944724221,"version":"3.53.1"},"reference-count":100,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,5,28]],"date-time":"2026-05-28T00:00:00Z","timestamp":1779926400000},"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                    Phosphorus (P) is essential for plant growth, and its deficiency significantly limits crop productivity. Plant biostimulants offer a sustainable approach to enhance plant tolerance to nutrient stress and reduce fertilizer use. Among these, humic substances (HS) derived from leonardite show promise, though their physiological effects under P deficiency remain unclear. This study evaluated the impact of a leonardite-derived HS on lettuce (\n                    <jats:italic>Lactuca sativa<\/jats:italic>\n                    L.) grown under two P regimes: high P (HP, 1 mM) and low P (LP, 0.2 mM). HS were applied at two concentrations via root (R1: 0.40; R2: 0.60 mL L\n                    <jats:sup>-<\/jats:sup>\n                    \u00b9) or foliar (F1: 7.50; F2: 10.00 mL L\n                    <jats:sup>-<\/jats:sup>\n                    \u00b9) treatments. Phosphorus shortage reduced shoot biomass and chlorophyll content, but stimulated root growth, total phenolic content (TPC), and antioxidant capacity. HS improved shoot growth across both P levels, with R2 being most effective at HP. Under LP, both HS doses produced similar effects. The enhanced tolerance to P deficiency conferred by HS was associated with improved root biometric traits, enhanced P acquisition (PAE), P utilization (PUtE), and overall P use efficiency (PUE), along with elevated acid phosphatase activity, chlorophyll, glycine betaine, TPC, and antioxidant capacity. All HS treatments enhanced root elongation; notably, F2 treatment increased root volume and diameter under LP. Furthermore, HS application modulated the leaf phenolic profile: F2 enhanced the accumulation of specific phenolics, while root-applied HS increased TPC and antioxidant response under LP. In conclusion, this leonardite-derived biostimulant demonstrated potential to enhance plant performance under P-limited conditions, supporting its use for more sustainable nutrient management.\n                  <\/jats:p>","DOI":"10.3389\/fpls.2026.1809888","type":"journal-article","created":{"date-parts":[[2026,5,28]],"date-time":"2026-05-28T04:44:50Z","timestamp":1779943490000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Leonardite-derived humic substances enhance phosphorus use efficiency in lettuce through integrated physiological and metabolic reprogramming"],"prefix":"10.3389","volume":"17","author":[{"given":"Santiago","family":"Atero-Calvo","sequence":"first","affiliation":[{"name":"Department of Plant Physiology, University of Granada","place":["Granada, Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Francesco","family":"Magro","sequence":"additional","affiliation":[{"name":"Sofbey S.A.","place":["Mendrisio, Switzerland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Giacomo","family":"Masetti","sequence":"additional","affiliation":[{"name":"Sofbey S.A.","place":["Mendrisio, Switzerland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Silvia","family":"Celletti","sequence":"additional","affiliation":[{"name":"Department of Agricultural, Forest and Food Sciences (DISAFA), University of Turin","place":["Grugliasco, Turin, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrea","family":"Ertani","sequence":"additional","affiliation":[{"name":"Department of Agricultural, Forest and Food Sciences (DISAFA), University of Turin","place":["Grugliasco, Turin, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Giuseppe","family":"Mannino","sequence":"additional","affiliation":[{"name":"Department of Life Sciences and Systems Biology, University of Turin","place":["Turin, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cinzia M.","family":"Bertea","sequence":"additional","affiliation":[{"name":"Department of Life Sciences and Systems Biology, University of Turin","place":["Turin, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Juan M.","family":"Ruiz","sequence":"additional","affiliation":[{"name":"Department of Plant Physiology, University of Granada","place":["Granada, Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michela","family":"Schiavon","sequence":"additional","affiliation":[{"name":"Department of Agricultural, Forest and Food Sciences (DISAFA), University of Turin","place":["Grugliasco, Turin, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,5,28]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.3390\/AGRONOMY14091939","article-title":"Reduced effect of commercial leonardite and seaweed extract on lettuce growth under mineral, organic, and no fertilization regimes","volume":"14","author":"Aguiar","year":"2024","journal-title":"Agronomy"},{"key":"B2","doi-asserted-by":"publisher","DOI":"10.3390\/agriculture10050147","article-title":"The effect of leonardite-derived amendments on soil microbiome structure and potato yield","volume":"10","author":"Akimbekov","year":"2020","journal-title":"Agriculture"},{"key":"B3","doi-asserted-by":"publisher","first-page":"244","DOI":"10.1016\/j.scienta.2018.03.047","article-title":"Ameliorative effects of calcium nitrate and humic acid on the growth, yield component and biochemical attribute of pepper (Capsicum annuum) plants grown under salt stress","volume":"236","author":"Akladious","year":"2018","journal-title":"Sci. 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