{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T04:26:31Z","timestamp":1787027191663,"version":"build-2736575974"},"reference-count":36,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2025,8,25]],"date-time":"2025-08-25T00:00:00Z","timestamp":1756080000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000921","name":"European Cooperation in Science and Technology","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100000921","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Plant Sci."],"abstract":"<jats:p>\n                    The Balkan Peninsula is a European biodiversity hotspot, home to 6,500 native vascular plant species, many of which are endemic. The region has diverse range of climates and complex topography, creating conditions that suit many woody ornamental, fruit, and forest species. Nevertheless, climate change, habitat destruction, invasive species, plant diseases, and agricultural practices threaten natural ecosystems and cultivated species. Many Balkan countries have addressed these challenges using advanced biotechnological approaches, including micropropagation,\n                    <jats:italic>in vitro<\/jats:italic>\n                    conservation, and\n                    <jats:italic>in vitro<\/jats:italic>\n                    selection for stress-tolerant genotypes. This paper provides a comprehensive overview of\n                    <jats:italic>in vitro<\/jats:italic>\n                    plant biotechnology progress in the Balkan countries from the beginning of the 21\n                    <jats:sup>st<\/jats:sup>\n                    century to the present, with a focus on woody horticultural and forest species. The primary objectives of scientific research include optimizing media composition for all components and micropropagation stages, as well as effective initial explant selection. In addition, temporary immersion bioreactors, synthetic seed technology, and cryopreservation techniques have been explored to enhance plant production and conservation. Scientific research for woody ornamentals and fruit tree species has progressed in Albania, Bulgaria, Croatia, Greece, and Serbia while remaining limited in other Balkan countries. Forest tree production research is currently conducted in Bulgaria, Greece, and Serbia, with labs focusing on micropropagation and\n                    <jats:italic>ex situ<\/jats:italic>\n                    conservation. In addition to advances in scientific research, several commercial companies operate in the Balkans, with establishments in Serbia, Greece, Bulgaria, and North Macedonia, that produce\n                    <jats:italic>in vitro<\/jats:italic>\n                    -derived planting material for fruit trees and woody ornamental plants. Despite this progress, research in the Balkan countries remains fragmented, emphasizing the need to strengthen regional collaboration and knowledge exchange to promote agricultural development processes and biotechnological applications in this region. This review represents the first exploration of this topic in the Balkans and successfully unites researchers from several countries. It highlights key scientific advances in\n                    <jats:italic>in vitro<\/jats:italic>\n                    biotechnology for woody plants, identifies challenges, and proposes solutions. The value of interdisciplinary collaboration is emphasized through the optimization of\n                    <jats:italic>in vitro<\/jats:italic>\n                    methodologies, the promotion of germplasm conservation, and the sustainable use of plant genetic resources in the Balkans.\n                  <\/jats:p>","DOI":"10.3389\/fpls.2025.1586013","type":"journal-article","created":{"date-parts":[[2025,8,25]],"date-time":"2025-08-25T05:27:12Z","timestamp":1756099632000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":8,"title":["Unveiling the Balkans\u2019 advances: In vitro biotechnology of woody plants in the early 21st century"],"prefix":"10.3389","volume":"16","author":[{"given":"Valbona","family":"Sota","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lilyana","family":"Nacheva","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dejan","family":"Bo\u0161njak","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eleni","family":"Abraham","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sla\u0111ana","family":"Jevremovi\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Branislav","family":"Cvjetkovi\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Vladislava","family":"Galovi\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Darko","family":"Jevremovi\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zvjezdana","family":"Markovi\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Efigjeni","family":"Kongjika","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sanja","family":"Bogunovi\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Svjetlana","family":"Zeljkovi\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Vlatko","family":"Andonovski","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Vanja","family":"Dani\u010di\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tatjana","family":"Vujovi\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2025,8,25]]},"reference":[{"key":"B1","article-title":"'Balkans'","author":"Allcock","year":"2024","journal-title":"Encyclopedia Britannica"},{"key":"B2","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1080\/13102818.1995.10818849","article-title":"Bulgarian plant biotechnology","volume":"9","author":"Atanassov","year":"1995","journal-title":"Biotechnol. Biotechnol. Equip."},{"key":"B3","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1080\/13102818.2002.10819178","article-title":"Challenges in front of the Bulgarian plant biotechnology","volume":"16","author":"Atanassov","year":"2002","journal-title":"Biotechnol. Biotechnol. Equip."},{"key":"B4","volume-title":"Biodiversity conservation and sustainable forestry","author":"Dave","year":"2003"},{"key":"B5","doi-asserted-by":"publisher","first-page":"1144","DOI":"10.1016\/j.tplants.2023.05.012","article-title":"CRISPR\/Cas-mediated plant genome editing: outstanding challenges a decade after implementation","author":"Cardi","year":"2023","journal-title":"Trends Plant Sci."},{"key":"B6","unstructured":"Skopje, Macedonia\n          CBD\n          CBD: \n          \n          2022"},{"key":"B7","doi-asserted-by":"publisher","first-page":"301","DOI":"10.3372\/wi.46.46303","article-title":"Vascular plants of Greece: An annotated checklist. Supplement","volume":"46","author":"Dimopoulos","year":"2016","journal-title":"Willdenowia"},{"key":"B8","unstructured":"European Environment Agency: The European reporting framework for adaptation\n          \n          2023"},{"key":"B9","unstructured":"Europe plant tissue culture market. Historic data: 2020-2021; forecast period: 2023-2028\n          \n          2022"},{"key":"B10","doi-asserted-by":"publisher","first-page":"141","DOI":"10.2298\/GSF14S1141G","article-title":"Nove biotehnologije u \u0161umarstvu Srbije","author":"Galovi\u0107","year":"2014","journal-title":"Glasnik \u0161umarskog fakulteta (Specialno izdanje)"},{"key":"B11","doi-asserted-by":"crossref","first-page":"104","DOI":"10.3832\/ifor0500-002","article-title":"Workshop COST E52 \u201cEvaluation of beech genetic resources for sustainable forestry","volume":"2","author":"Giannini","year":"2009","journal-title":"iForest"},{"key":"B12","article-title":"Biodiversity and nature conservation in the Western Balkans \u2013 civil society, (local) politics, international actors, and the media in dialogue","author":"Hansj\u00f6rg","year":"2022","journal-title":"S\u00fcdosteuropa Mitt."},{"key":"B13","article-title":"\u201cEffective acclimatization of in vitro cultured plants: methods, physiology and genetics,\u201d","author":"Hazarika","year":"2006","journal-title":"Floriculture, Ornamental and Plant Biotechnology: Advances and Topical Issues, Volume II."},{"key":"B14","first-page":"37","article-title":"Metodat e bioteknologjise \"in vitro\" per fitimin e bimeve te painfektuara","volume":"10","author":"Kongjika","year":"1995","journal-title":"Punime te Institutit t\u00eb K\u00ebrkimeve Biologjike"},{"key":"B15","doi-asserted-by":"publisher","first-page":"153","DOI":"10.15177\/seefor.20-18","article-title":"In vitro lead tolerance testing in white poplar genotypes on acidic medium","volume":"11","author":"Kova\u010devi\u0107","year":"2020","journal-title":"South-East Eur. Forestry"},{"key":"B16","unstructured":"Lovri\u0107\n              M.\n            \n            \n              Lovri\u0107\n              N.\n            \n          \n          Integration of nature protection in forest policy in Croatia. INTEGRATE Country Report for Croatia. European Forest Institute, EFICEEC \u2013 EFISEE Regional Office\n          \n          2013"},{"key":"B17","first-page":"327","article-title":"Collecting plant genetic resources in Europe: A survey of legal requirements and practical experiences","volume-title":"Implementing the nagoya protocol","author":"Maggioni","year":"2017"},{"key":"B18","first-page":"125","article-title":"Raznovidnost i poteklo na florata na republika makedonija","volume-title":"Opening adresses, contributions and bibliography of the new members of the Macedonian Academy of sciences and Arts","author":"Matevski","year":"2013"},{"key":"B19","volume-title":"The Red Book of vascular flora of the Republic of Kosovo","author":"Millaku","year":"2013"},{"key":"B20","volume-title":"Fourth national report of Montenegro to the convention on biological diversity","author":"Bu\u0161kovi\u0107","year":"2010"},{"key":"B21","article-title":"Impacts of climate change on agriculture and recommendations for adaptation measures in the Western Balkans. (GFA Consulting Group on behalf of the Federal Ministry of Food and Agriculture (BMEL)","author":"M\u00fcller","year":"2022"},{"key":"B22","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1002\/9781119607045.ch27","article-title":"Invasive alien species in the Balkan Peninsula","volume-title":"Invasive Alien Species: Observations and Issues from Around the World","author":"Panjkovi\u0107","year":"2021"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.5586\/asbp.914","article-title":"Application and Improvement of in vitro culture systems for commercial production of ornamental, fruit, and industrial plants in Poland","volume":"91","author":"Podwyszy\u0144ska","year":"2022","journal-title":"Acta Societatis Botanicorum Poloniae"},{"key":"B24","volume-title":"Flowers of Greece and the balkans-A field guide","author":"Polunin","year":"1980"},{"key":"B25","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1016\/S0044-328X(75)80123-1","article-title":"Embryogenesis in tissue culture of Corylus avellana L. Z","volume":"77","author":"Radojevi\u0107","year":"1975","journal-title":"Phlanzen Physiol."},{"key":"B26","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1007\/978-1-4020-2854-0_2","article-title":"The physical geography of the Balkans and nomenclature of place names","volume-title":"Balkan Biodiversity: Pattern and Process in the European Hotspot","author":"Reed","year":"2004"},{"key":"B27","doi-asserted-by":"publisher","first-page":"51","DOI":"10.2298\/GSF14S1051S","article-title":"Forest genetic resources in Serbia - state and recommendations for improvement in this area","author":"\u0160ija\u010di\u0107-Nikoli\u0107","year":"2014","journal-title":"Bull. Faculty Forestry"},{"key":"B28","article-title":"Key to the native and foreign vascular plants in Bulgaria","volume-title":"Interactive extended and supplemented edition","author":"Stoyanov","year":"2022"},{"key":"B29","volume-title":"The Plant-Life of the Balkan Peninsula - a Phytogeographical Study","author":"Turill","year":"1929"},{"key":"B30","doi-asserted-by":"publisher","first-page":"896","DOI":"10.1002\/elsc.201900041","article-title":"Use of bioreactor systems in the propagation of forest trees","volume":"19","author":"Vidal","year":"2019","journal-title":"Eng. Life Sci."},{"key":"B31","volume-title":"The way back: brain drain and prosperity in the Western Balkans","author":"Vracic","year":"2018"},{"key":"B32","doi-asserted-by":"publisher","first-page":"5","DOI":"10.5937\/topola2006005","article-title":"Effect of IBA and TIBA on rhizogenesis of Wild cherry in vitro","volume":"206","author":"Vuksanovi\u0107","year":"2020","journal-title":"Topola"},{"key":"B33","first-page":"13","article-title":"The influence of drought on growth and development of white poplar shoots in vitro","author":"Vuksanovi\u0107","year":"2019","journal-title":"Topola\/Poplar"},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.3390\/life13030780","article-title":"Tissue culture\u2014a sustainable approach to explore plant stresses","volume":"13","author":"Wijerathna-Yapa","year":"2023","journal-title":"Life"},{"key":"B35","volume-title":"Cutting-edge technologies in the agriculture of Plovdiv district","author":"Yordanov","year":"1986"},{"key":"B36","volume-title":"National biodiversity strategy of Albania, (2012-2020)","author":"Zeneli","year":"2014"}],"container-title":["Frontiers in Plant Science"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2025.1586013\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,8,25]],"date-time":"2025-08-25T05:27:17Z","timestamp":1756099637000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2025.1586013\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,8,25]]},"references-count":36,"alternative-id":["10.3389\/fpls.2025.1586013"],"URL":"https:\/\/doi.org\/10.3389\/fpls.2025.1586013","relation":{},"ISSN":["1664-462X"],"issn-type":[{"value":"1664-462X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,8,25]]},"article-number":"1586013"}}