Impact of anthropogenic disturbance on tree non-spatial structure, seasonal physiology and nutrient dynamics in urban forests of Pinus koraiensis

Abstract

Urban forest ecosystems are the key contributors to the green spaces within metropolitan landscapes, offering numerous ecological services. But the combined effects of anthropogenic activities driven by population growth and escalating resource demands have precipitated rapid modification in forest structure, extent, and environmental conditions (i.e., nutrient dynamics) of such areas. This study investigates how human disturbances affect Korean pine urban forests through examinations of non-spatial structure and physiological, biochemical and soil responses. Tree height decreased (R² = 0.463, p < 0.001) while canopy cover increased (R² = 0.132, p < 0.001) as disturbance intensity intensified. Needle indole-3-acetic acid (IAA) and gibberellic acid (GA₃) content varied significantly (p = 0.017 and p = 0.049) across the various disturbance levels, with significant seasonal cytokinin and GA₃ changes under very low disturbance (p = 0.023 and 0.040) and pronounced IAA and GA₃ under heavy disturbance (p = 0.026 and 0.024). Needle moisture content varies significantly (p = 0.029) with increasing disturbance and significant seasonal changes only when disturbances reached heavy intensity level (p = 0.0138), and chlorophyll (Chl) a (p = 0.0002), Chl b (p = 0.009), and total Chl (p = 0.0004) were disturbance sensitive. In addition, annual glucose (p < 0.001), starch (p < 0.01), and non-structural carbohydrate (p < 0.001) content in needle, branches, and root were exhibited significant change with increasing disturbance, seasonal variations especially found in heavily disturbed sites (p < 0.05). Nutrient element content (i.e., carbon and nitrogen ratio, total phosphorus, and total potassium) in needles, branches, roots, and soil showed significant differences (p < 0.05) with increasing disturbance intensities, except for phosphorus in roots which was not significant. These findings quantified the threshold at which urban disturbances impair Korean pine resilience and point out the urgency of regulating anthropogenic disturbances and restoring soil nutrient conditions. This study advances urban forest ecology by linking disturbance intensity to physiological thresholds and nutrient depletion, supporting for integrated conservation frameworks that prioritize urban forests alongside protected areas. Future research should target urban Korean pine growth, particularly regarding height increments, and highlights the importance of nutrient management and disturbance mitigation.

Keywords

Urban forest / Anthropogenic disturbance / Forest structure / Nutrients dynamics / Seasonal physiology / Korean pine

References

[1] Aryal PC, Aryal C, Bhusal K, Chapagain D, Dhamala MK, Maharjan SR, Chhetri PK. Forest structure and anthropogenic disturbances regulate plant invasion in urban forests. Urban Ecosyst, 2022, 25(2): 367-377

[2] Ashraf M, Harris PJC. Photosynthesis under stressful environments: an overview. Photosynthetica, 2013, 51(2): 163-190

[3] Ashton MS, Kelty MJThe practice of silviculture: applied forest ecology, 201810th editionUKJohn Wiley & Sons Ltd

[4] Bacelar E, Pinto T, Anjos R, Morais MC, Oliveira I, Vilela A, Cosme F. Impacts of climate change and mitigation strategies for some abiotic and biotic constraints influencing fruit growth and quality. Plants, 2024, 13(14): Article ID: 1942

[5] Bahadur Budha P, Rai P, Katel P, Khadka A. Vulnerability assessment of peoples exposed to landslides in Panchase of Nepal using analytical hierarchy process. Int J Environ, 2020, 9(2): 81-103

[6] Bao Y, Liu X, Feng CH, Niu MX, Liu C, Wang HL, Yin WL, Xia XL. Light and light signals regulate growth and development in woody plants. Forests, 2024

[7] Barnes BV, Xü Z, Zhao SD. Forest ecosystems in an old-growth pine–mixed hardwood forest of the Changbai Shan preserve in northeastern China. Can J for Res, 1992, 22(2): 144-160

[8] Battisti C, Poeta G, Fanelli G. An introduction to disturbance ecology: a road map for wildlife management and conservation. Springer International Publishing, 2016

[9] Beninde J, Veith M, Hochkirch A. Biodiversity in cities needs space: a meta-analysis of factors determining intra-urban biodiversity variation. Ecol Lett, 2015, 18(6): 581-592

[10] Boggs J, Sun G, Jones D, McNulty SG. Effect of soils on water quantity and quality in Piedmont forested headwater watersheds of North Carolina. JAWRA J Am Water Resour Assoc, 2013, 49(1): 132-150

[11] Brant AN, Chen HYH. Patterns and mechanisms of nutrient resorption in plants. Crit Rev Plant Sci, 2015, 34(5): 471-486

[12] Burgess TI, Oliva J, Sapsford SJ, Sakalidis ML, Balocchi F, Paap T. Anthropogenic disturbances and the emergence of native diseases: a threat to forest health. Curr for Rep, 2022, 8(2): 111-123

[13] Burton PJ, Jentsch A, Walker LR. The ecology of disturbance interactions. Bioscience, 2020, 70(10): 854-870

[14] Caron S, Garvey SM, Gewirtzman J, Schultz K, Bhatnagar JM, Driscoll C, Hutyra LR, Templer pH. Urbanization and fragmentation have opposing effects on soil nitrogen availability in temperate forest ecosystems. Glob Change Biol, 2023, 29(8): 2156-2171

[15] Castro-Camba R, Sánchez C, Vidal N, Vielba JM. Interactions of gibberellins with phytohormones and their role in stress responses. Horticulturae, 2022

[16] Chen YN, Chen YP, Zhou HH, Hao XM, Zhu CG, Fu AH, Yang YH, Li WH. Research advances in plant physiology and ecology of desert riparian forests under drought stress. Forests, 2022

[17] Chen X, Xiao KT, Deng RX, Wu L, Cui LJ, Ning H, Ai XR, Chen H. Projecting the future redistribution of Pinus koraiensis (Pinaceae: Pinoideae: Pinus) in China using machine learning. Front for Glob Change, 2024, 7 Article ID: 1326319

[18] Cheng SK, Xu ZR, Su Y, Zhen L. Spatial and temporal flows of China’s forest resources: development of a framework for evaluating resource efficiency. Ecol Econ, 2010, 69(7): 1405-1415

[19] Cornelissen JHC, Lavorel S, Garnier E, Díaz S, Buchmann N, Gurvich DE, Reich PB, ter Steege H, Morgan HD, van der Heijden MGA, Pausas JG, Poorter H. A handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Aust J Bot, 2003, 51(4): 335-380

[20] Crudo J. Effects of soil compaction and organic matter removal on selected soil properties and tree growth in a boreal ecosystem of northeastern British Columbia. Uni Bri Col, 2024

[21] Czaja M, Kołton A, Muras P. The complex issue of urban trees—stress factor accumulation and ecological service possibilities. Forests, 2020

[22] da Vila Gonçalves  (2023) Metabolic shifts associated with drought and drought-like symptoms induced by pine pitch canker disease. Univ De Lis (Portugal)

[23] De Diego N, Pérez-Alfocea F, Cantero E, Lacuesta M, Moncaleán P. Physiological response to drought in Radiata pine: phytohormone implication at leaf level. Tree Physiol, 2012, 32(4): 435-449

[24] Doucet TC, Duinker PN, Zurba M, Steenberg JWN, Charles JD. Perspectives of successes and challenges in collaborations between non-governmental organization and local government on urban forest management. Urban for Urban Green, 2024, 93 Article ID: 128220

[25] Dovletyarova EA, Mosina LV, Vasenev VI, Ananyeva ND, Patlseva A, Ivashchenko KV. Monitoring and assessing anthropogenic influence on soil’s health in urban forests: the case from Moscow City. Adaptive soil management: from theory to practices, 2017SingaporeSpringer531-557

[26] Dror I, Yaron B, Berkowitz B. The human impact on all soil-forming factors during the anthropocene. ACS Environ Au, 2022, 2(1): 11-19

[27] Dunne T, Western D, Dietrich WE. Effects of cattle trampling on vegetation, infiltration, and erosion in a tropical rangeland. J Arid Environ, 2011, 75(1): 58-69

[28] EL Sabagh A, Islam MS, Hossain A, Iqbal MA, Mubeen M, Waleed M, Reginato M, Battaglia M, Ahmed S, Rehman A, Arif M, Athar HUR, Ratnasekera D, Danish S, Ali Raza M, Rajendran K, Mushtaq M, Skalicky M, Brestic M, Soufan W, Fahad S, Pandey S, Kamran M, Datta R, Abdelhamid MT. Phytohormones as growth regulators during abiotic stress tolerance in plants. Front Agron, 2022, 4: 765068

[29] Emiru N, Gebrekidan H. Effect of land use changes and soil depth on soil organic matter, total nitrogen and available phosphorus contents of soils in Senbat Watershed, Western Ethiopia. ARPN J Agri Bio Sci, 2013, 8(3): 206-212

[30] Esperon-Rodriguez M, Rymer PD, Power SA, Barton DN, Cariñanos P, Dobbs C, Eleuterio AA, Escobedo FJ, Hauer R, Hermy M, Jahani A, Onyekwelu JC, Östberg J, Pataki D, Randrup TB, Rasmussen T, Roman LA, Russo A, Shackleton C, Solfjeld I, van Doorn NS, Wells MJ, Wiström B, Yan PB, Yang J, Tjoelker MG. Assessing climate risk to support urban forests in a changing climate. Plants People Planet, 2022, 4(3): 201-213

[31] Feng G, Svenning JC, Mi XC, Jia Q, Rao MD, Ren HB, Bebber DP, Ma KP. Anthropogenic disturbance shapes phylogenetic and functional tree community structure in a subtropical forest. For Ecol Manag, 2014, 313: 188-198

[32] Fornal-Pieniak B, Ollik M, Schwerk A. Impact of different levels of anthropogenic pressure on the plant species composition in woodland sites. Urban for Urban Green, 2019, 38: 295-304

[33] Forner A, Valladares F, Bonal D, Granier A, Grossiord C, Aranda I. Extreme droughts affecting Mediterranean tree species’ growth and water-use efficiency: the importance of timing. Tree Physiol, 2018, 38(8): 1127-1137

[34] Frelich LE, Reich PB. Minireviews: neighborhood effects, disturbance severity, and community stability in forests. Ecosystems, 1999, 2(2): 151-166

[35] Frolking S, Palace MW, Clark DB, Chambers JQ, Shugart HH, Hurtt GC. Forest disturbance and recovery: a general review in the context of spaceborne remote sensing of impacts on aboveground biomass and canopy structure. J Geophys Res Biogeosci, 2009, 114(G2): 2008JG000911

[36] Furze ME, Wainwright DK, Huggett BA, Knipfer T, McElrone AJ, Brodersen CR. Ecologically driven selection of nonstructural carbohydrate storage in oak trees. New Phytol, 2021, 232(2): 567-578

[37] Gangwar S, Singh VP, Tripathi DK, Chauhan DK, Prasad SM, Maurya JN (2014) Plant responses to metal stress. In: Emerging technologies and management of crop stress tolerance. Elsevier, pp 215–248. https://doi.org/10.1016/b978-0-12-800875-1.00010-7

[38] Gierth M, Mäser P. Potassium transporters in plants–involvement in K+ acquisition, redistribution and homeostasis. FEBS Lett, 2007, 581(12): 2348-2356

[39] Gorgens EB, Nunes MH, Jackson T, Coomes D, Keller M, Reis CR, Valbuena R, Rosette J, de Almeida DRA, Gimenez B, Cantinho R, Motta AZ, Assis M, de Souza Pereira FR, Spanner G, Higuchi N, Ometto JP. Resource availability and disturbance shape maximum tree height across the Amazon. Glob Chang Biol, 2021, 27(1): 177-189

[40] Greene CS, Millward AA. Getting closure: the role of urban forest canopy density in moderating summer surface temperatures in a large city. Ecosystems, 2017, 20(1): 141-156

[41] Grieve IC. Human impacts on soil properties and their implications for the sensitivity of soil systems in Scotland. CATENA, 2001, 42(2–4): 361-374

[42] Haas J. Ecosystem services from space as evaluation metric of human well-being in deprived urban areas of the majority world. Urban inequalities from space, 2024ChamSpringer International Publishing259-285

[43] He M, Zhou GY, Yuan TF, van Groenigen KJ, Shao JJ, Zhou XH. Grazing intensity significantly changes the C: N: P stoichiometry in grassland ecosystems. Ecosystems, 2020, 29(2): 355-369

[44] Huang L, Ning ZY, Zhang XL. Impacts of caterpillar disturbance on forest net primary production estimation in China. Ecol Indic, 2010, 10(6): 1144-1151

[45] Ji YR, Zhang P, Shen HL. Competition intensity affects growing season nutrient dynamics in Korean pine trees and their microhabitat soil in mixed forest. For Ecol Manag, 2023, 539 Article ID: 121018

[46] Jim CY. Soil compaction as a constraint to tree growth in tropical & subtropical urban habitats. Envir Conserv, 1993, 20(1): 35-49

[47] Jones JA, Wei XH, Archer E, Bishop K, Blanco JA, Ellison D, Gush MB, McNulty SG, van Noordwijk M, Creed IF. Forest-water interactions under global change. Forest-water interactions, 2020ChamSpringer International Publishing589-624

[48] Khan A, Karim MR, Mohammed KMG, Sinha K, Sultana F, Mukul SA, Arfin-Khan MAS. Anthropogenic disturbance modifies tree functional traits in the only remnant swamp forest of Bangladesh. Front Ecol Evol, 2023, 11: 1062764

[49] Khan F, Siddique AB, Shabala S, Zhou MX, Zhao CC. Phosphorus plays key roles in regulating plants’ physiological responses to abiotic stresses. Plants, 2023, 12(15): Article ID: 2861

[50] Konijnendijk CC. Evidence-based guidelines for greener, healthier, more resilient neighbourhoods: introducing the 3–30–300 rule. J for Res, 2023, 34(3): 821-830

[51] Kozlowski TT. Soil compaction and growth of woody plants. Scand J for Res, 1999, 14(6): 596-619

[52] Le Maitre DC, Gush MB, Dzikiti S. Impacts of invading alien plant species on water flows at stand and catchment scales. AoB Plants, 2015, 7 Article ID: plv043

[53] Lebedev AV. Changes in the growth of Scots pine (Pinus sylvestris L.) stands in an urban environment in European Russia since 1862. J for Res, 2023, 34(5): 1279-1287

[54] Leduc N, Roman H, Barbier F, Péron T, Huché-Thélier L, Lothier J, Demotes-Mainard S, Sakr S. Light signaling in bud outgrowth and branching in plants. Plants, 2014, 3(2): 223-250

[55] Li Z, Zhang ZW, Wang YJ, Wang PC, Xu YR, Zhou ZX. Influence of anthropogenic disturbances on understory plant diversity of urban forests in Wuhan, Central China. Sai Mal, 2012, 41(12): 1495-1501

[56] Li Y, He NP, Hou JH, Xu L, Liu CC, Zhang JH, Wang QF, Zhang XM, Wu XQ. Factors influencing leaf chlorophyll content in natural forests at the biome scale. Front Ecol Evol, 2018, 6 Article ID: 64

[57] Li T, Zheng WW, Zhou ZJ, Zhang SR, Xu XX, Pu YL, Li H. Soil phosphorus variation regulated by changes in land use spatial patterns during urbanization in western Chengdu, China. Glob Ecol Conserv, 2021, 27 Article ID: e01576

[58] Li B, Ma X, Saha S, Wu HB, Zhang P, Shen HL. Korean pines demonstrate cold resilience through non-structural carbohydrate concentrations despite light deprivation during the growing season. Forests, 2023

[59] Li LJ, Zheng QF, Jiang W, Xiao NY, Zeng FR, Chen G, Mak M, Chen ZH, Deng FL. Molecular regulation and evolution of cytokinin signaling in plant abiotic stresses. Plant Cell Physiol, 2022, 63(12): 1787-1805

[60] Li WK, Li B, Ma X, Saha S, Wu HB, Zhang P, Shen HL. Physiological and biochemical traits of needles imply that understory light conditions in the growing season may be favorable to Pinus koraiensis trees. Forests, 2023

[61] Li XY, Bleisch WV, Hu WQ, Li Q, Wang HJ, Chen ZZ, Bai R, Jiang XL. Human disturbance increases spatiotemporal associations among mountain forest terrestrial mammal species. Elife, 2024, 12 Article ID: RP92457

[62] Li P, Wang ZJ, Ran K. Coexistence of multiple leaf nutrient resorption strategies in urban forests. Glob Ecol Conserv, 2025, 57 Article ID: e03367

[63] Liang DC, Huang GL. Influence of urban tree traits on their ecosystem services: a literature review. Land, 2023

[64] Liang SY, Tan T, Wu DZ, Li CJ, Jing HQ, Wu JW. Seasonal variations in carbon, nitrogen, and phosphorus of Pinus yunnanenis at different stand ages. Front Plant Sci, 2023, 14 Article ID: 1107961

[65] Lim SD, Mayer JA, Yim WC, Cushman JC. Plant tissue succulence engineering improves water-use efficiency, water-deficit stress attenuation and salinity tolerance in Arabidopsis. Plant J, 2020, 103(3): 1049-1072

[66] Liu JY, Sherif SM. Hormonal orchestration of bud dormancy cycle in deciduous woody perennials. Front Plant Sci, 2019, 10: 1136

[67] Liu RS, Wang DM. C: N: P stoichiometric characteristics and seasonal dynamics of leaf-root-litter-soil in plantations on the Loess Plateau. Ecol Indic, 2021, 127 Article ID: 107772

[68] Liu WD, Su JR, Li SF, Lang XD, Huang XB. Non-structural carbohydrates regulated by season and species in the subtropical monsoon broad-leaved evergreen forest of Yunnan Province, China. Sci Rep, 2018, 8(1): Article ID: 1083

[69] Liu GC, Xing YJ, Wang QG, Wang L, Feng Y, Yin ZW, Wang XC, Liu T. Long-term nitrogen addition regulates root nutrient capture and leaf nutrient resorption in Larix gmelinii in a boreal forest. Eur J for Res, 2021, 140(4): 763-776

[70] Liu JY, Lu SZ, Liu CC, Hou DJ. Nutrient reallocation between stem and leaf drives grazed grassland degradation in Inner Mongolia, China. BMC Plant Biol, 2022, 22(1): Article ID: 505

[71] Lv HL, Wang WJ, He XY, Xiao L, Zhou W, Zhang B. Quantifying tree and soil carbon stocks in a temperate urban forest in Northeast China. Forests, 2016

[72] Mandre M, Lukjanova A. Biochemical and structural characteristics of Scots pine (Pinus sylvestris L.) in an alkaline environment. Est J Ecol, 2011, 60(4): 264-283

[73] Mao QG, Chen H, Gurmesa GA, Gundersen P, Ellsworth DS, Gilliam FS, Wang C, Zhu F, Ye Q, Mo JM, Lu XK. Negative effects of long-term phosphorus additions on understory plants in a primary tropical forest. Sci Total Environ, 2021, 798 Article ID: 149306

[74] Martin-Benito D, Pederson N, Lanter C, Köse N, Doğan M, Bugmann H, Bigler C. Disturbances and climate drive structure, stability, and growth in mixed temperate old-growth rainforests in the Caucasus. Ecosystems, 2020, 23(6): 1170-1185

[75] Maynard DG, Paré D, Thiffault E, Lafleur B, Hogg KE, Kishchuk B. How do natural disturbances and human activities affect soils and tree nutrition and growth in the Canadian boreal forest?. Environ Rev, 2014, 22(2): 161-178

[76] McIntyre S, Lavorel S, Landsberg J, Forbes TDA. Disturbance response in vegetation–towards a global perspective on functional traits. J Veg Sci, 1999, 10(5): 621-630

[77] Mensah S, Houehanou TD, Sogbohossou EA, Assogbadjo AE, Glèlè Kakaï R. Effect of human disturbance and climatic variability on the population structure of Afzelia africana Sm. ex pers. (Fabaceae–Caesalpinioideae) at country broad-scale (Bénin, West Africa). S Afr N J Bot, 2014, 95: 165-173

[78] Mikalajunas M, Pretzsch H, Mozgeris G, Linkevičius E, Augustaitiene I, Augustaitis A. Scots pine’s capacity to adapt to climate change in hemi-boreal forests in relation to dominating tree increment and site condition. iForest, 2021, 14(5): 473-482

[79] Mikhailova TA, Afanasieva LV, Kalugina OV, Shergina OV, Taranenko EN. Changes in nutrition and pigment complex in pine (Pinus sylvestris L.) needles under technogenic pollution in Irkutsk region, Russia. J for Res, 2017, 22(6): 386-392

[80] Moles AT, Perkins SE, Laffan SW, Flores-Moreno H, Awasthy M, Tindall ML, Sack L, Pitman A, Kattge J, Aarssen LW, Anand M, Bahn M, Blonder B, Cavender-Bares J, Cornelissen JHC, Cornwell WK, Díaz S, Dickie JB, Freschet GT, Griffiths JG, Gutierrez AG, Hemmings FA, Hickler T, Hitchcock TD, Keighery M, Kleyer M, Kurokawa H, Leishman MR, Liu K, Niinemets Ü, Onipchenko V, Onoda Y, Penuelas J, Pillar VD, Reich PB, Shiodera S, Siefert A, Sosinski EEJr, Soudzilovskaia NA, Swaine EK, Swenson NG, van Bodegom PM, Warman L, Weiher E, Wright IJ, Zhang HX, Zobel M, Bonser SP. Which is a better predictor of plant traits: temperature or precipitation?. J Veg Sci, 2014, 25(5): 1167-1180

[81] Monaco P, Baldoni A, Naclerio G, Scippa GS, Bucci A. Impact of plant-microbe interactions with a focus on poorly investigated urban ecosystems-a review. Microorganisms, 2024, 12(7): Article ID: 1276

[82] Montague M (2021) Linking competition and nonstructural carbon dynamics in american chestnut (Castanea dentata). Purdue University, 30504704

[83] Mukherjee D. Nutrient and its management Prospect and challenges under the changing environment scenario. Advances in plant physiology-an international treatise series, 2014Jodhpur, IndiaScientific Publishers413-442

[84] Muneer MA, Afridi MS, Abu Bakar Saddique M, Chen XH, Zaib-Un-Nisa YXJ, Farooq I, Munir MZ, Yang WH, Ji BM, Zheng CY, Wu LQ. Nutrient stress signals: elucidating morphological, physiological, and molecular responses of fruit trees to macronutrients deficiency and their management strategies. Sci Hortic, 2024, 329: 112985

[85] Munné-Bosch S, Alegre L. Changes in carotenoids, tocopherols and diterpenes during drought and recovery, and the biological significance of chlorophyll loss in Rosmarinus officinalis plants. Planta, 2000, 210(6): 925-931

[86] Muramatsu S, Kojima K, Igasaki T, Azumi Y, Shinohara K. Inhibition of the light-independent synthesis of chlorophyll in pine cotyledons at low temperature. Plant Cell Physiol, 2001, 42(8): 868-872

[87] Muthukkaruppan E, Lavanya AK, Chinnathambi V, Suku AT, Paul S (2024) Application of bioinoculants in horticulture, plantation, and forest farming: is it truly ecologically sustainable? . In: Bio-inoculants in horticultural crops. Elsevier, pp 21–48. https://doi.org/10.1016/b978-0-323-96005-2.00003-9

[88] Nandal A, Rani S, Yadav SS, Kaushik N, Kataria N, Hasanpuri P, Lal R. Soil quality under different tree species in an urban university campus: a multidimensional study. Environ Earth Sci, 2024, 83(21): Article ID: 613

[89] Nielsen KS, Marteau TM, Bauer JM, Bradbury RB, Broad S, Burgess G, Burgman M, Byerly H, Clayton S, Espelosin D, Ferraro PJ, Fisher B, Garnett EE, Jones JPG, Otieno M, Polasky S, Ricketts TH, Trevelyan R, van der Linden S, Veríssimo D, Balmford A. Biodiversity conservation as a promising frontier for behavioural science. Nat Hum Behav, 2021, 5(5): 550-556

[90] Noh NJ, Son Y, Lee SK, Yoon TK, Seo KW, Kim C, Lee WK, Bae SW, Hwang J. Influence of stand density on soil CO₂ efflux for a Pinus densiflora forest in Korea. J Plant Res, 2010, 123(4): 411-419

[91] Onaindia M, Dominguez I, Albizu I, Garbisu C, Amezaga I. Vegetation diversity and vertical structure as indicators of forest disturbance. For Ecol Manag, 2004, 195(3): 341-354

[92] Owens PN. Soil erosion and sediment dynamics in the Anthropocene: a review of human impacts during a period of rapid global environmental change. J Soils Sediments, 2020, 20(12): 4115-4143

[93] Pak U, Guo QX, Liu ZL, Wang XG, Liu YK, Jin GZ. Spatial distribution of Pinus koraiensis trees and community-level spatial associations in broad-leaved Korean pine mixed forests in northeastern China. Plants, 2023, 12(16): 2906

[94] Pandey BK, Bennett MJ. Uncovering root compaction response mechanisms: new insights and opportunities. J Exp Bot, 2024, 75(2): 578-583

[95] Park A, Puettmann K, Wilson E, Messier C, Kames S, Dhar A. Can boreal and temperate forest management be adapted to the uncertainties of 21st century climate change?. Crit Rev Plant Sci, 2014, 33(4): 251-285

[96] Peterson CJ, Pickett STA. Forest reorganization: a case study in an old-growth forest catastrophic blowdown. Ecology, 1995, 76(3): 763-774

[97] Petrova ST, Yurukova LD, Velcheva IG. Assessment of the urban trees health status on the base of nutrient and pigment content in their leaves. J Bio Sci Bio, 2014, 3(1): 69-77

[98] Pickett STA, Cadenasso ML. Altered resources, disturbance, and heterogeneity: a framework for comparing urban and non-urban soils. Urban Ecosyst, 2009, 12(1): 23-44

[99] Piper FI, Paula S. The role of nonstructural carbohydrates storage in forest resilience under climate change. Curr for Rep, 2020, 6(1): 1-13

[100] Polinko AD, Willis JL, Sharma A, Guldin JM. Stand-level structural characteristics dictate hurricane resistance and resilience more than silvicultural regime in longleaf pine woodlands. For Ecol Manag, 2022, 526 Article ID: 120585

[101] Prietzel J, Stetter U, Klemmt HJ, Rehfuess KE. Recent carbon and nitrogen accumulation and acidification in soils of two Scots pine ecosystems in Southern Germany. Plant Soil, 2006, 289(1): 153-170

[102] Qin RF, Zhao QR, Gu CR, Wang C, Zhang L, Zhang HG. Analysis of oxidase activity and transcriptomic changes related to cutting propagation of hybrid larch. Sci Rep, 2023, 13(1): Article ID: 1354

[103] Rangel Pinagé E, Bell DM, Longo M, Gao SC, Keller M, Silva CA, Ometto JP, Köhler P, Frankenberg C, Huete A. Forest structure and solar-induced fluorescence across intact and degraded forests in the Amazon. Remote Sens Environ, 2022, 274 Article ID: 112998

[104] Reich PB. The world-wide ‘fast–slow’ plant economics spectrum: a traits manifesto. J Ecol, 2014, 102(2): 275-301

[105] Ren ZB, Zhao HB, Fu Y, Xiao L, Dong YL. Effects of urban street trees on human thermal comfort and physiological indices: a case study in Changchun City, China. J for Res, 2022, 33(3): 911-922

[106] Ren XX, Huang C, Zhu ZZ, Wang L, Wang N, Mu XY, Luo DR, Zhang J, Xiang LX, Huang XZ. Anthropogenic forest degradation and its impact on soil erosion in the mountainous region of northern China. CATENA, 2024, 243 Article ID: 108217

[107] Sakib N, Poudel TR, Hao YQ, Roberts NJ, Iddrisu AQ, Adhikari S, Zhang P. Effects of different thinning intensities on carbon storage in Pinus koraiensis middle-aged plantations in Northeast China. Forests, 2024

[108] Santner A, Calderon-Villalobos LIA, Estelle M. Plant hormones are versatile chemical regulators of plant growth. Nat Chem Biol, 2009, 5(5): 301-307

[109] Sardans J, Grau O, Chen HYH, Janssens IA, Ciais P, Piao SL, Peñuelas J. Changes in nutrient concentrations of leaves and roots in response to global change factors. Glob Change Biol, 2017, 23(9): 3849-3856

[110] Seidl R, Senf C. Changes in planned and unplanned canopy openings are linked in Europe’s forests. Nat Commun, 2024, 15(1): Article ID: 4741

[111] Seidl R, Turner MG. Post-disturbance reorganization of forest ecosystems in a changing world. Proc Natl Acad Sci USA, 2022, 119(28): Article ID: e2202190119

[112] Shah SH, Islam S, Mohammad F, Siddiqui MH. Gibberellic acid: a versatile regulator of plant growth, development and stress responses. J Plant Growth Regul, 2023, 42(12): 7352-7373

[113] Shanahan DF, Fuller RA, Bush R, Lin BB, Gaston KJ. The health benefits of urban nature: how much do we need?. Bioscience, 2015, 65(5): 476-485

[114] Shao PS, Han HY, Sun JK, Xie HT. Effects of global change and human disturbance on soil carbon cycling in boreal forest: a review. Pedosphere, 2023, 33(1): 194-211

[115] Shi F, Meng QL, Pan L, Wang JS. Root damage of street trees in urban environments: an overview of its hazards, causes, and prevention and control measures. Sci Total Environ, 2023, 904 Article ID: 166728

[116] Sieghardt M, Mursch-Radlgruber E, Paoletti E, Couenberg E, Dimitrakopoulus A, Rego F, Hatzistathis A, Randrup TB. The abiotic urban environment: impact of urban growing conditions on urban vegetation. Urban forests and trees, 2005Berlin, HeidelbergSpringer281-323

[117] Singh H. Climate change and environmental pollution: biological consequences on urban forests. Urban forests, climate change and environmental pollution, 2024ChamSpringer Nature Switzerland1-21

[118] Singh R, Chavan SB, Kakade VD, Morade AS, Singh AK, Rawale GB, Uthappa AR, Keerthika A, Chichaghare AR, Gurav S, Reddy KS. Ecosystem services provided by urban and peri-urban forests. Urban forests, climate change and environmental pollution, 2024ChamSpringer Nature Switzerland417-445

[119] Skalak J, Nicolas KL, Vankova R, Hejatko J. Signal integration in plant abiotic stress responses via multistep phosphorelay signaling. Front Plant Sci, 2021, 12 Article ID: 644823

[120] Smith P, House JI, Bustamante M, Sobocká J, Harper R, Pan GX, West PC, Clark JM, Adhya T, Rumpel C, Paustian K, Kuikman P, Cotrufo MF, Elliott JA, McDowell R, Griffiths RI, Asakawa S, Bondeau A, Jain AK, Meersmans J, Pugh TAM. Global change pressures on soils from land use and management. Glob Change Biol, 2016, 22(3): 1008-1028

[121] Song Y, Li XH, Zhang MY, Xiong C. Spatial specificity of metabolism regulation of abscisic acid-imposed seed germination inhibition in Korean pine (Pinus koraiensis Sieb et Zucc). Front Plant Sci, 2024, 15 Article ID: 1417632

[122] Sørensen LH, Mikola J, Kytöviita MM, Olofsson J. Trampling and spatial heterogeneity explain decomposer abundances in a sub-Arctic grassland subjected to simulated reindeer grazing. Ecosystems, 2009, 12(5): 830-842

[123] Sun WX, Li QQ, Qiao B, Jia KT, Li CY, Zhao CJ. Advances in plant–soil feedback driven by root exudates in forest ecosystems. Forests, 2024

[124] Swoczyna T, Kalaji HM, Bussotti F, Mojski J, Pollastrini M. Environmental stress—what can we learn from chlorophyll a fluorescence analysis in woody plants? A review. Front Plant Sci, 2022, 13 Article ID: 1048582

[125] Takahashi M, Feng ZZ, Mikhailova TA, Kalugina OV, Shergina OV, Afanasieva LV, Heng RKJ, Majid NMA, Sase H. Air pollution monitoring and tree and forest decline in East Asia: a review. Sci Total Environ, 2020, 742 Article ID: 140288

[126] Tang LN, Shao GF, Piao ZJ, Dai LM, Jenkins MA, Wang SX, Wu G, Wu JG, Zhao JZ. Forest degradation deepens around and within protected areas in East Asia. Biol Conserv, 2010, 143(5): 1295-1298

[127] Thakrey M, Singh L, Jhariya MK, Tomar A, Singh AK, Toppo S. Impact of disturbance on biomass, carbon, and nitrogen storage in vegetation and on soil properties of tropical dry deciduous forest in Chhattisgarh, India. Land Degrad Dev, 2022, 33(11): 1810-1820

[128] Tian JF, Wang BY, Zhang CR, Li WD, Wang SJ. Mechanism of regional land use transition in underdeveloped areas of China: a case study of Northeast China. Land Use Policy, 2020, 94 Article ID: 104538

[129] Tuomi M, Väisänen M, Ylänne H, Brearley FQ, Barrio IC, Bråthen AK, Eischeid I, Forbes BC, Jónsdóttir IS, Kolstad AL, Macek P, Petit Bon M, Speed JDM, Stark S, Svavarsdóttir K, Thórsson J, Bueno CG. Stomping in silence: conceptualizing trampling effects on soils in polar tundra. Funct Ecol, 2021, 35(2): 306-317

[130] Tuzhilkina VV. Response of the pigment system of conifers to long-term industrial air pollution. Russ J Ecol, 2009, 40(4): 227-232

[131] Vishal B, Kumar PP. Regulation of seed germination and abiotic stresses by gibberellins and abscisic acid. Front Plant Sci, 2018, 9 Article ID: 838

[132] Wallace B, Bulmer C, Hope G, Curran M, Philpott T, Murray M. Soil compaction and organic matter removal effects on soil properties and tree growth in the Interior Douglas-fir zone of southern British Columbia. For Ecol Manage, 2021, 494 Article ID: 119268

[133] Wang AY, Han SJ, Zhang JH, Wang M, Yin XH, Fang LD, Yang D, Hao GY. The interaction between nonstructural carbohydrate reserves and xylem hydraulics in Korean pine trees across an altitudinal gradient. Tree Physiol, 2018, 38(12): 1792-1804

[134] Wang MZ, Li HL, Liu CX, Dong BC, Yu FH. Adaptive plasticity in response to light and nutrient availability in the clonal plant Duchesnea indica. J Plant Ecol, 2022, 15(4): 795-807

[135] Wang K, She DQ, Zhang XT, Wang YY, Wen H, Yu JH, Wang QG, Han SJ, Wang WJ. Tree richness increased biomass carbon sequestration and ecosystem stability of temperate forests in China: interacted factors and implications. J Environ Manage, 2024, 368 Article ID: 122214

[136] Wang YY, Dai XZ, Chen XL, Zhang D, Lin GQ, Zhou YH, Wang TY, Cui YL. Effects of urbanization and forest type on species composition and diversity, forest characteristics, biomass carbon sink, and their associations in Changchun, Northeast China: implications for urban carbon stock improvement. J for Res, 2024, 35(1): Article ID: 36

[137] Ward EB, Doroski DA, Felson AJ, Hallett RA, Oldfield EE, Kuebbing SE, Bradford MA. Positive long-term impacts of restoration on soils in an experimental urban forest. Ecol Appl, 2021, 31(5): Article ID: e02336

[138] Watson GW, Hewitt AM, Custic M, Lo M. The management of tree root systems in urban and suburban settings II: a review of strategies to mitigate human impacts. Isa, 2014, 40(5): 249-271

[139] Wolters H, Jürgens G. Survival of the flexible: hormonal growth control and adaptation in plant development. Nat Rev Genet, 2009, 10(5): 305-317

[140] Woś B, Chodak M, Józefowska A, Pietrzykowski M. Influence of tree species on carbon, nitrogen, and phosphorus stocks and stoichiometry under different soil regeneration scenarios on reclaimed and afforested mine and post-fire forest sites. Geoderma, 2022, 415 Article ID: 115782

[141] Xiao L, Wang WJ, He XY, Lv HL, Wei CH, Zhou W, Zhang B. Urban-rural and temporal differences of woody plants and bird species in Harbin City, northeastern China. Urban for Urban Green, 2016, 20: 20-31

[142] Xu HY, Wang H, Prentice IC, Harrison SP. Leaf carbon and nitrogen stoichiometric variation along environmental gradients. Biogeosciences, 2023, 20(22): 4511-4525

[143] Xu J, Lei L, Zeng LX, Ni YY, Jian ZJ, Deng XX, Xiao WF. The responses of C allocation of new needle and fine root affected the phosphorus adaptation of Pinus massoniana seedlings. J Soil Sci Plant Nutr, 2024, 24(1): 295-307

[144] Xu ZZ, Qin LH, Zhou G, SiQing B, Du WX, Meng SW, Yu J, Sun Z, Liu QJ. Exploring carbon sequestration in broad-leaved Korean pine forests: insights into photosynthetic and respiratory processes. Sci Total Environ, 2024, 906 Article ID: 167421

[145] Yadav A, Bohra NK, Singh H. Forest ecosystem adaptation to climate change: morphological and physiological responses. Forests and climate change, 2024SingaporeSpringer Nature269-289

[146] Yang J, Zhang J, Wang Z, Zhu Q, Wang W. Hormonal changes in the grains of rice subjected to water stress during grain filling. Plant Physiol, 2001, 127(1): 315-323

[147] Zhang PQ, Liu YJ, Chen X, Yang Z, Zhu MH, Li YP. Pollution resistance assessment of existing landscape plants on Beijing streets based on air pollution tolerance index method. Ecotoxicol Environ Saf, 2016, 132: 212-223

[148] Zhang Y, Li C, Wang ML. Linkages of C: N: P stoichiometry between soil and leaf and their response to climatic factors along altitudinal gradients. J Soils Sediments, 2019, 19(4): 1820-1829

[149] Zhang PP, Yin MZ, Zhang XJ, Wang QT, Wang RH, Yin HJ. Differential aboveground-belowground adaptive strategies to alleviate N addition-induced P deficiency in two Alpine coniferous forests. Sci Total Environ, 2022, 849 Article ID: 157906

[150] Zhang PP, Ding JX, Wang QT, McDowell NG, Kong DL, Tong YD, Yin HJ. Contrasting coordination of non-structural carbohydrates with leaf and root economic strategies of Alpine coniferous forests. New Phytol, 2024, 243(2): 580-590

[151] Zhao PJ, Yuan DD. Population growth and urban–rural structure. Population growth and sustainable transport in China, 2023SingaporeSpringer Nature59-96

[152] Zheng LL, Zhao Q, Yu ZY, Zhao SY, Zeng DH. Altered leaf functional traits by nitrogen addition in a nutrient-poor pine plantation: a consequence of decreased phosphorus availability. Sci Rep, 2017, 7(1): Article ID: 7415

[153] Zhou GY, Zhou XH, He YH, Shao JJ, Hu ZH, Liu RQ, Zhou HM, Hosseinibai S. Grazing intensity significantly affects belowground carbon and nitrogen cycling in grassland ecosystems: a meta-analysis. Glob Change Biol, 2017, 23(3): 1167-1179

[154] Zhou G, Liu QJ, Xu ZZ, Du WX, Yu J, Meng SW, Zhou H, Qin LH, Shah S. How can the shade intolerant Korean pine survive under dense deciduous canopy?. For Ecol Manag, 2020, 457 Article ID: 117735

[155] Zhou XY, Kutchartt E, Hernández J, Corvalán P, Promis Á, Zwanzig M. Determination of optimal tree height models and calibration designs for Araucaria araucana and Nothofagus pumilio in mixed stands affected to different levels by anthropogenic disturbance in South-Central Chile. Ann for Sci, 2023, 80(1): Article ID: 18

[156] Zhu LJ, Zhang J, Camarero JJ, Cooper DJ, Cherubini P, Yuan DY, Wang XC. Drivers and spatiotemporal patterns of post-drought growth resilience of four temperate broad-leaved trees. Agric for Meteor, 2023, 342 Article ID: 109741

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