<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD JATS (Z39.96) Journal Archiving and Interchange DTD with OASIS Tables with MathML3 v1.4 20241031//EN" "https://jats.nlm.nih.gov/archiving/1.4/JATS-archive-oasis-article1-4-mathml3.dtd">
<article xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:ali="http://www.niso.org/schemas/ali/1.0/" dtd-version="1.4" article-type="research-article" xml:lang="en"><front><journal-meta><journal-title-group><journal-title xml:lang="ru">Теоретическая и прикладная экология</journal-title></journal-title-group><issn publication-format="print">1995-4301</issn><issn publication-format="electronic">2618-8406</issn></journal-meta><article-meta><article-id pub-id-type="doi">10.25750/1995-4301-2026-1-056-064</article-id><article-categories><subj-group><subject>Other</subject></subj-group></article-categories><title-group><article-title xml:lang="ru">Сезонные изменения флуоресценции хлорофилла для оценки стрессоустойчивости сосны сибирской на верхнем пределе распространения</article-title><trans-title-group xml:lang="en"><trans-title>Seasonal changes in chlorophyll fluorescence to assess the stress tolerance of Siberian pines at treeline</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author"><name-alternatives><name xml:lang="ru"><surname>Пахарькова</surname><given-names>Н. В.</given-names></name><name xml:lang="en"><surname>Pakharkova</surname><given-names>N. V.</given-names></name></name-alternatives><xref ref-type="aff" rid="aff1"/><xref ref-type="aff" rid="aff2"/><email>nina.pakharkova@yandex.ru</email><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-0377-3263</contrib-id></contrib><contrib contrib-type="author"><name-alternatives><name xml:lang="ru"><surname>Гетте</surname><given-names>И. Г.</given-names></name><name xml:lang="en"><surname>Gette</surname><given-names>I. G.</given-names></name></name-alternatives><xref ref-type="aff" rid="aff1"/><xref ref-type="aff" rid="aff2"/><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-7372-1499</contrib-id></contrib><contrib contrib-type="author"><name-alternatives><name xml:lang="ru"><surname>Гаевский</surname><given-names>Н. А.</given-names></name><name xml:lang="en"><surname>Gaevsky</surname><given-names>N. A.</given-names></name></name-alternatives><xref ref-type="aff" rid="aff1"/><xref ref-type="aff" rid="aff2"/><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-3747-9858</contrib-id></contrib><contrib contrib-type="author"><name-alternatives><name xml:lang="ru"><surname>Рудиковская</surname><given-names>Е. Г.</given-names></name><name xml:lang="en"><surname>Rudikovskaya</surname><given-names>E. G.</given-names></name></name-alternatives><xref ref-type="aff" rid="aff3"/><xref ref-type="aff" rid="aff4"/><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-5709-2370</contrib-id></contrib><contrib contrib-type="author"><name-alternatives><name xml:lang="ru"><surname>Рудиковский</surname><given-names>А. В.</given-names></name><name xml:lang="en"><surname>Rudikovskii</surname><given-names>A. V.</given-names></name></name-alternatives><xref ref-type="aff" rid="aff3"/><xref ref-type="aff" rid="aff4"/><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-8567-0318</contrib-id></contrib><contrib contrib-type="author"><name-alternatives><name xml:lang="ru"><surname>Масенцова</surname><given-names>И. В.</given-names></name><name xml:lang="en"><surname>Masentsova</surname><given-names>I. V.</given-names></name></name-alternatives><xref ref-type="aff" rid="aff1"/><xref ref-type="aff" rid="aff2"/><contrib-id contrib-id-type="orcid">https://orcid.org/0009-0000-4038-6583</contrib-id></contrib><contrib contrib-type="author"><name-alternatives><name xml:lang="ru"><surname>Калабина</surname><given-names>А. А.</given-names></name><name xml:lang="en"><surname>Kalabina</surname><given-names>A. A.</given-names></name></name-alternatives><xref ref-type="aff" rid="aff1"/><xref ref-type="aff" rid="aff2"/><contrib-id contrib-id-type="orcid">https://orcid.org/0009-0000-3563-2460</contrib-id></contrib><contrib contrib-type="author"><name-alternatives><name xml:lang="ru"><surname>Гавриков</surname><given-names>В. Л.</given-names></name><name xml:lang="en"><surname>Gavrikov</surname><given-names>V. L.</given-names></name></name-alternatives><xref ref-type="aff" rid="aff1"/><xref ref-type="aff" rid="aff2"/><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-7816-0536</contrib-id></contrib><aff-alternatives id="aff1"><aff><institution xml:lang="en">Siberian Federal University (Krasnoyarsk, Russia)</institution></aff></aff-alternatives><aff-alternatives id="aff2"><aff><institution xml:lang="ru">Сибирский федеральный университет (Красноярск, Россия)</institution></aff></aff-alternatives><aff-alternatives id="aff3"><aff><institution xml:lang="en">Siberian Institute of Plant Physiology and Biochemistry of the Siberian Branch of the Russian Academy of Sciences (Irkutsk, Russia)</institution></aff></aff-alternatives><aff-alternatives id="aff4"><aff><institution xml:lang="ru">Сибирский институт физиологии и биохимии растений (Иркутск, Россия)</institution></aff></aff-alternatives></contrib-group><pub-date pub-type="epub" iso-8601-date="2026-03-25"><day>25</day><month>03</month><year>2026</year></pub-date><issue>1</issue><fpage>56</fpage><lpage>64</lpage><self-uri xlink:href="http://www.envjournal.ru/ari/v2026/v1/26106.pdf" xlink:title="http://www.envjournal.ru/ari/v2026/v1/26106.pdf">http://www.envjournal.ru/ari/v2026/v1/26106.pdf</self-uri><self-uri content-type="pdf" xlink:href="publication-b8ec1c1e-31c4-4b1c-afe4-390a0841309d.pdf" xlink:title="PDF"/><abstract xml:lang="ru"><p>Из-за изменения климата границы ареалов многих растений смещаются как в широтном, так и в высотном направлениях, в частности, граница лесов в горах смещается вверх. На склоне хребта Ергаки (Западный Саян) была заложена трансекта с четырьмя пробными площадями, пересекающая верхнюю границу леса при переходе от горно-таёжного к гольцово-тундровому поясу. Выявлено, что растения сосны сибирской (Pinus sibirica Du Tour) используют различные стратегии выживания при распространении выше границы леса, что проявляется в изменении формы кроны со стволовой на стланниковую, а также в сезонных изменениях пигментного комплекса и параметров флуоресценции хлорофилла хвои. Во всех местообитаниях P. sibirica важным параметром для прогнозирования увеличения массы хвои, которое, в свою очередь, обеспечивает продуктивность всего растения, является максимальная скорость нециклического переноса электронов в летнее время. Весной и осенью (при переходе от состояния зимнего покоя к вегетации и обратно) в пигментном комплексе хвои уменьшается соотношение хлорофиллов и каротиноидов от лесных к горно-тундровым фитоценозам. Экземпляры сосны сибирской, растущие выше границы леса и имеющие механизмы рассеивания избыточной световой энергии путём светоиндуцированного нефотохимического тушения, имеют явное преимущество в наборе массы хвои перед другими растениями с тех же пробных площадей. Для деревьев с пробных площадей, расположенных в лесных фитоценозах, высокая доля нефотохимического тушения является скорее помехой. Рассмотренные параметры флуоресценции хлорофилла могут быть использованы для выявления особей P. sibirica, устойчивых к условиям обитания выше границы леса, и их последующего использования в селекции для увеличения площади лесных массивов в условиях изменения климата.</p></abstract><abstract xml:lang="en" abstract-type="summary"><p>Due to climate change, the boundaries of the ranges of many plants are shifting both in latitudinal and altitudinal directions. In particular, the forest boundary shifts upward in the mountains. A transect with four sampling plots was laid on the slope of the Ergaki Ridge (Western Sayan), crossing the treeline (upper border) of the forest as it transits from the mountain taiga to the rocky-tundra belt. It has been revealed that Siberian pine (Pinus sibirica Du Tour) plants use various survival strategies when spreading above the treeline, which manifests itself in a change in the shape of the crown form from stem-like to krummholz, as well as seasonal changes in the pigment complex and parameters of chlorophyll fluorescence in needles. In all habitats of P. sibirica, an important parameter for predicting an increase in the needle weight, which, in turn, ensures the productivity of the entire plant, is the maximum rate of non-cyclic electron transfer in the summer. In spring and autumn (in the course of transition from winter dormancy to vegetation and back), the ratio of chlorophylls and carotenoids from forest to mountain-tundra communities decreases in the pigment complex of needles. Siberian pine specimens growing above the treeline and having mechanisms for dissipating excess light energy by light-induced non-photochemical fluorescence quenching have a clear advantage in gaining needle weight over other plants from the same sample plots. For trees from sample plots located in forest environment, a high proportion of nonphotochemical fluorescence quenching is rather a disadvantage. The considered parameters of chlorophyll fluorescence can be used to identify P. sibirica individuals resistant to habitat conditions above the treeline and their subsequent use in breeding to increase the area of woodlands under climate change.</p></abstract><kwd-group xml:lang="ru"><kwd>Pinus sibirica</kwd><kwd>верхняя граница леса</kwd><kwd>флуоресценция</kwd><kwd>хлорофилл</kwd><kwd>каротиноиды</kwd></kwd-group><kwd-group xml:lang="en"><kwd>Pinus sibirica</kwd><kwd>treeline</kwd><kwd>fluorescence</kwd><kwd>chlorophyll</kwd><kwd>carotenoids</kwd></kwd-group><funding-group><funding-statement xml:lang="en">This research was funded by Russian Science Foundation to the research project: “Intrapopulation variability of ecophysiological features of Siberian pine (Pinus sibirica Du Tour) trees under climate change”, grant number 23-24-00251.</funding-statement></funding-group></article-meta></front><back><ref-list><ref id="ref1"><mixed-citation publication-type="other" xml:lang="en">1. Lyu L., Suvanto S., Nöjd P., Henttonen H.M., Mäkinen H., Zhang Q.B. Tree growth and its climate signal along latitudinal and altitudinal gradients: comparison of tree rings between Finland and the Tibetan Plateau // Biogeosciences. 2017. V. 14. P. 3083–3095. doi: 10.5194/bg-14-3083-2017</mixed-citation></ref><ref id="ref2"><mixed-citation publication-type="other" xml:lang="en">2. Klinge M., Dulamsuren C., Erasmi S., Karger D.N., Hauck M. Climate effects on vegetation vitality at the treeline of boreal forests of Mongolia // Biogeosciences. 2018. V. 15. P. 1319–1333. doi: 10.5194/bg-15-1319-2018</mixed-citation></ref><ref id="ref3"><mixed-citation publication-type="other" xml:lang="en">3. Kruse S., Gerdes A., Kath N.J., Epp L.S., Stoof-Leichsenring K.R., Pestryakova L.A., Herzschuh U. Dispersal distances and migration rates at the arctic treeline in Siberia – a genetic and simulation-based study // Biogeosciences. 2019. V. 16. No. 6. P. 1211–1224. doi: 10.5194/bg-16-1211-2019</mixed-citation></ref><ref id="ref4"><mixed-citation publication-type="other" xml:lang="en">4. Kharuk V.I., Im S.T., Petrov I.A. Alpine ecotone in the Siberian Mountains: vegetation response to warming // J. Mt. Sci. 2021. V. 18. P. 3099–3108. doi: 10.1007/s11629-021-6876-2</mixed-citation></ref><ref id="ref5"><mixed-citation publication-type="other" xml:lang="en">5. Grigoriev A.A., Shalaumova Yu.V., Vyukhin S.O., Balakin D.S., Kukarskikh V.V., Vyukhina A.A., Camarero J.J., Moiseev P.A. Upward treeline shifts in two regions of Subarctic Russia are governed by summer thermal and winter snow conditions // Forests. 2022. V. 13. Article No. 174. doi: 10.3390/f13020174</mixed-citation></ref><ref id="ref6"><mixed-citation publication-type="other" xml:lang="en">6. Moiseev P.A., Hagedor F., Balakin D.S., Bubnov M.O., Devi N.M., Kukarskih V.V., Mazepa V.S., Viyukhin S.O., Viyukhina A.A., Grigoriev A.A. Stand biomass at treeline ecotone in Russian Subarctic Mountains is primarily related to species composition but its dynamics driven by improvement of climatic conditions // Forests. 2022 V. 13. Article No. 254. doi: 10.3390/f13020254</mixed-citation></ref><ref id="ref7"><mixed-citation publication-type="other" xml:lang="en">7. Petrov I.A., Shushpanov A.S., Golyukov A.S., Dvinskaya M.L., Kharuk V.I. Dynamics of tree and shrub vegetation in the Eastern Sayan Mountain Tundra // Russ. J. Ecol. 2021. V. 52. P. 399–405. doi: 10.1134/S1067413621050118</mixed-citation></ref><ref id="ref8"><mixed-citation publication-type="other" xml:lang="en">8. Kharuk V.I., Im S.T., Dvinskaya M.L. Forest-tundra ecotone response to climate change in the Western Sayan Mountains, Siberia // Scand. J. For. Res. 2010. V. 25. No. 3. P. 224–233. doi: 10.1080/02827581003766959</mixed-citation></ref><ref id="ref9"><mixed-citation publication-type="other" xml:lang="en">9. Petrov I.A., Kharuk V.I., Dvinskaya M.L., Im S.T. Reaction of coniferous trees in the Kuznetsk Alatau alpine forest-tundra ecotone to climate change // Contemp. Probl. Ecol. 2015. V. 8. P. 423–430. doi: 10.1134/S1995425515040137</mixed-citation></ref><ref id="ref10"><mixed-citation publication-type="other" xml:lang="en">10. Pakharkova N., Borisova I., Sharafutdinov R., Gavrikov V. Photosynthetic pigments in Siberian pine and fir under climate warming and shift of the timberline // Forests. 2020. V. 11. Article No. 63. doi: 10.3390/f11010063</mixed-citation></ref><ref id="ref11"><mixed-citation publication-type="other" xml:lang="en">11. Tantsyrev N.V., Shobairi S.O.R., Usoltsev V.A., Sun L., Zhang H., Li C., He J., Hemmati Roudbari S., Asghari Beirami B., Ayombekov Q. Geographical peculiarities of Pinus sibirica Du Tour natural regeneration as related to its seed crops // Curr. World Environ. 2024. V. 19 No. 1. P. 109–123. doi: 10.12944/CWE.19.1.4</mixed-citation></ref><ref id="ref12"><mixed-citation publication-type="other" xml:lang="en">12. Velisevich S.N., Bender O.G., Goroshkevich S.N The influence of scion donor tree age on the growth and morphogenesis of Siberian stone pine grafts // New Forests. 2020. V. 52. P. 473–491. doi: 10.1007/s11056-020-09805-2</mixed-citation></ref><ref id="ref13"><mixed-citation publication-type="other" xml:lang="en">13. Furbank R.T., Tester M. Phenomics – technologies to relieve the phenotyping bottleneck // Trends Plant Sci. 2011. V. 16. No. 12. P. 635–644. doi: 10.1016/j.tplants.2011.09.005</mixed-citation></ref><ref id="ref14"><mixed-citation publication-type="other" xml:lang="en">14. Demidchik V.V., Shashko A.Y., Bandarenka U.Y., Smolikova G.N., Przhevalskaya D.A., Charnysh M.A., Pozhvanov G.A., Barkosvkyi A.V., Smolich I.I., Sokolik A.I., Yu M., Medvedev S.S. Plant phenomics: fundamental bases, software and hardware platforms, and machine learning // Russ. J. Plant Physiol. 2020. V. 67. No. 3. P. 397–412. doi: 10.1134/S1021443720030061</mixed-citation></ref><ref id="ref15"><mixed-citation publication-type="other" xml:lang="en">15. Rolfe S.A., Scholes J.D. Chlorophyll fluorescence imaging of plant-pathogen interactions // Protoplasma. 2010. V. 247. P. 163–175. doi: 10.1007/s00709-010-0203-z</mixed-citation></ref><ref id="ref16"><mixed-citation publication-type="other" xml:lang="en">16. Dutta S., Cruz J.A., Jiao Y., Chen J., Kramer D.M., Osteryoung K.W. Non-invasive, whole-plant imaging of chloroplast movement and chlorophyll fluorescence reveals photosynthetic phenotypes independent of chloroplast photorelocation defects in chloroplast division mutants // Plant J. 2015. V. 84. No. 2. P. 428–442. doi: 10.1111/tpj.13009</mixed-citation></ref><ref id="ref17"><mixed-citation publication-type="other" xml:lang="en">17. Kalaji H.M., Jajoo A., Oukarroum A., Brestic M., Zivcak M., Samborska I.A., Cetner M.D., Łukasik I., Goltsev V., Ladle R.J. Chlorophyll a fluorescence as a tool to monitor physiological status of plants under abiotic stress conditions // Acta Physiol. Plant. 2016. V. 38. Article No. 102. doi: 10.1007/s11738-016-2113-y</mixed-citation></ref><ref id="ref18"><mixed-citation publication-type="other" xml:lang="en">18. Smolikova G., Kreslavski V., Shiroglazova O., Bilova T., Sharova E., Frolov A., Medvedev S. Photochemical activity changes accompanying the embryogenesis of pea (Pisum sativum) with yellow and green cotyle-dons // Funct. Plant Biol. 2018. V. 45. No. 2. P. 228–235. doi: 10.1071/FP16379</mixed-citation></ref><ref id="ref19"><mixed-citation publication-type="other" xml:lang="en">19. Pakharkova N.V., Gaevsky N.A., Gette I.G. Parameters of Pinus sibirica photosynthetic apparatus for phenotyping trees in the high-altitude zone of the Western Sayan // J. Sib. Fed. Univ. Biol. 2024. V. 17. No. 2. P. 119–133 (in Russian).</mixed-citation></ref><ref id="ref20"><mixed-citation publication-type="other" xml:lang="en">20. WALZ – High Quality Instrumentation for Plant Sciences [Internet resource] http://www.walz.com (Accessed: 15.11.2024).</mixed-citation></ref><ref id="ref21"><mixed-citation publication-type="other" xml:lang="en">21. Rudikovskii A.V., Stolbicova A.V., Rudikovskaya E.G., Dudareva L.V. Role of phytohormones in the formation of dwarf and tall Siberian crabapple (Malus bаccata L. Borkh.) // Zemdirbyste. 2019. V. 106. No. 2. P. 167–172. doi: 10.13080/z-a.2019.106.022</mixed-citation></ref><ref id="ref22"><mixed-citation publication-type="other" xml:lang="en">22. Lichtenthaler H.K. Chlorophylls and carotenoids: Pigments of photosynthetic biomembranes // Methods in Enzymology / Eds. S.P. Colowick, N.O. Kaplan. San Diego: Academic Press, 1987. P. 350–382. doi: 10.1016/0076-6879(87)48036-1</mixed-citation></ref><ref id="ref23"><mixed-citation publication-type="other" xml:lang="en">23. Küpper H., Seibert S., Parameswaran A. Fast, sensitive, and inexpensive alternative to analytical pigment HPLC: quantification of chlorophylls and carotenoids in crude extracts by fitting with Gauss peak spectra // Anal. Chem. 2007. V. 79. No. 20. P. 7611–7627. doi: 10.1021/ac070236m</mixed-citation></ref><ref id="ref24"><mixed-citation publication-type="other" xml:lang="en">24. Rudikovskaya Е.G., Dudareva L.V., Shishparenok А.A., Osipova S.V., Stavitskaya Z.O., Rudikovskii А.V. Peculiarities of accumulation of photoprotective compounds by fruit of Siberian crabapple, Malus domestica Borkh. and their hybrids growing in conditions of Eastern Siberia // Sci. Hortic. 2019. V. 253. P. 303–308. doi: 10.1016/j.scienta.2019.04.034</mixed-citation></ref><ref id="ref25"><mixed-citation publication-type="other" xml:lang="en">25. Robakowski P. Susceptibility to low-temperature photoinhibition in three conifers differing in successional status // Tree Physiol. 2005. V. 25. No. 9. P. 1151–1160. doi: 10.1093/treephys/25.9.1151</mixed-citation></ref><ref id="ref26"><mixed-citation publication-type="other" xml:lang="en">26. Tuzhilkina V.V. Photosynthetic activity of coniferous undergrowth in connection with industrial logging // Theoretical and Applied Ecology. 2024. No. 2. P. 22–28 (in Russian). doi: 10.25750/1995-4301-2024-2-022-028</mixed-citation></ref><ref id="ref27"><mixed-citation publication-type="other" xml:lang="en">27. Öquist G., Huner N.P.A. Photosynthesis of overwintering evergreen plants // Annu. Rev. Plant Biol. 2003. V. 54. P. 329–355. doi: 10.1146/annurev.arplant.54.072402.115741</mixed-citation></ref><ref id="ref28"><mixed-citation publication-type="other" xml:lang="en">28. Ruban A.V. Evolution under the sun: optimizing light harvesting in photosynthesis // J. Exp. Bot. 2015. V. 66. No. 1. P. 7–23. doi: 10.1093/jxb/eru400</mixed-citation></ref><ref id="ref29"><mixed-citation publication-type="other" xml:lang="en">29. Pospíšil P. Mechanisms of non-photochemical chlorophyll fluorescence quenching in higher plants // Photosynthetica. 1997. V. 34. P. 343–355. doi: 10.1023/A:1006803832366</mixed-citation></ref><ref id="ref30"><mixed-citation publication-type="other" xml:lang="en">30. Maslova T.G., Markovskaya E.F., Slemnev N.N. Functions of carotenoids in leaves of higher plants (review) // Biol. Bull. Rev. 2021. V. 11. No. 5. P. 476–487. doi: 10.1134/S2079086421050078</mixed-citation></ref></ref-list></back></article>
