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Testing phaeophytinization as an index of ozone stress in trees

Journal of Forestry Research 2023年第4期34卷 1167-1174页

作者： Evgenios Agathokleous Mitsutoshi Kitao Takayoshi Koike School of Applied Meteorology Nanjing University of Information Science and Technology(NUIST)Nanjing 210044People’s Republic of China Research Faculty of Agriculture Hokkaido UniversitySapporoHokkaido 060-8589Japan Hokkaido Research Center Forestry and Forest Products Research Institute(FFPRI)Sapporo 062-8516Japan

Ground-level ozone pollution is a menace for vegetation in the northern hemisphere,limiting photosynthetic pigments and suppressing photosynthesis in trees and other types of plants.Phaeophytinization is the process of converting chlorophylls into phaeophytins,for example by acidifi cation.Ozone is a highly oxidizing molecule and well known to degrade chlorophylls;however,the eff ect of ozone on phaeophytinization in leaves of higher plants is largely unknown.To reveal ozone eff ect on phaeophytinization and evaluate the potential of phaeophytinization as an index of ozone stress in trees,the absorbance at the optical density of 665 nm was measured before(OD_(665))and after(OD_(665a))acidifi cation in three independent experiments with nearly 30 conditions of ozone exposure.Both current ambient and elevated ozone widely aff ected phaeophytinization,as indicated by decreases or increases in the phaeophytinization quotient OD_(665)/OD_(665a).These eff ects were commonly moderate to large in magnitude and practically signifi cant,and occurred even in ozone-asymptomatic leaves.It emerges that the ozone eff ect on phaeophytinization is bimodal,likely depending on the intensity of ozone stress.These results indicate a promising feature of OD_(665)/OD_(665a)as a thorough index of ozone stress in the future,but further studies are needed to reveal the underlying biochemical mechanisms of the bimodal eff ect on phaeophytinization.

关键词： Biological stress Chemical eff ect Chlorophyll oxidation Environmental stimuli Phaeophytin

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Single and combined eff ects of fertilization,ectomycorrhizal inoculation,and drought on container-grown Japanese larch seedlings

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Journal of Forestry Research 2023年第4期34卷 1077-1094页

作者： Evgenios Agathokleous Mitsutoshi Kitao Masabumi Komatsu Yutaka Tamai Hisanori Harayama Takayoshi Koike Collaborative Innovation Center on Forecast and Evaluation of Meteorological Disasters(CIC-FEMD) Nanjing University of Information Science and TechnologyNanjing 210044JiangsuPeople’s Republic of China Research Center for Global Changes and Ecosystem Carbon Sequestration and Mitigation School of Applied MeteorologyNanjing University of Information Science and TechnologyNanjing 210044JiangsuPeople’s Republic of China Hokkaido Research Center Forestry and Forest Products Research Institute(FFPRI)Forest Research and Management Organization7 HitsujigaokaSapporoHokkaido 062-8516Japan Department of Mushroom Science and Forest Microbiology Forestry and Forest Products Research Institute(FFPRI)Matsunosato 1Tsukuba 305-8687Japan Research Faculty of Agriculture Hokkaido UniversityKita 9 Nishi 9SapporoHokkaido 060-8589Japan Department of Plant Ecology Forestry and Forest Products Research Institute(FFPRI)Matsunosato 1Tsukuba 305-8687Japan

Climate change can intensify drought in many regions of the world and lead to more frequent drought events or altered cycles of soil water status.Therefore,it is important to enhance the tolerance to drought and thus health,vigor,and success of transplantation seedlings used in the forestry by modifying fertilization and promoting mycorrhization.Here,we sowed seeds of Japanese larch(Larix kaempferi)in 0.2-L containers with 0.5 g(low fertilization;LF)or 2 g(high fertilization;HF)of slow-release fertilizer early in the growing season.One month later,we irrigated seedlings with non-sterilized ectomycorrhizal inoculum(ECM)or sterilized solution(non-ECM),and after about 2 months,plants were either kept well watered(WW;500 mL water/plant/week)or subjected to drought(DR;50 mL water per plant/week)until the end of the growing season.HF largely stimulated plant growth and above-and belowground biomass production,eff ects that are of practical signifi cance,but caused a small decrease in stomatal conductance(Gs 390)and transpiration rate(E 390),which in practice is insignifi cant.ECM treatment resulted in moderate inhibition of seedling growth and biomass and largely canceled out the enhancement of biomass and foliar K content by HF.DR caused a large decrease in CO 2 assimilation,and enhanced stomatal closure and induced senescence.DR also largely depleted foliar Mg and enriched foliar K.Although DR caused a large decrease in foliar P content in LF,it moderately increased P in HF.Likewise,DR increased foliar K in HF but not in LF,and decreased foliar P in ECM plants but not in non-ECM plants.Conversely,ECM plants exhibited a large enrichment in foliar P under WW and had a lower water potential under DR when grown in LF.These results indicate that the drought tolerance and health and vigor of Japanese larch seedlings can be modifi ed by soil fertility and soil microorganisms.This study provides a basis for new multifactorial research programs aimed at producing seedlings of superior quality for forestation under climate cha

关键词： Drought Ectomycorrhizae Fertilizer eff ect Oxidative stress Stressor interactions

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Ethylenediurea(EDU)effects on hybrid larch saplings exposed to ambient or elevated ozone over three growing seasons

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Journal of Forestry Research 2022年第1期33卷 117-135页

作者： Evgenios Agathokleous Mitsutoshi Kitao Takayoshi Koike Key Laboratory of Agrometeorology of Jiangsu Province School of Applied MeteorologyNanjing University of Information Science and Technology(NUIST)Nanjing 210044People’s Republic of China Research Faculty of Agriculture Hokkaido UniversitySapporoHokkaido 060-8589Japan Hokkaido Research Center Forestry and Forest Products Research Institute(FFPRI)Sapporo 062-8516Japan Research Center for Eco-Environmental Science Chinese Academy of ScienceBeijing 100085People’s Republic of China

Ground-level ozone(O_(3))pollution is a persistent environmental issue that can lead to adverse effects on trees and wood production,thus indicating a need for forestry interventions to mediate O_(3) effects.We treated hybrid larch(Larix gmelinii var.japonica×L.kaempferi)saplings grown in nutrient-poor soils with 0 or 400 mg L^(-1) water solutions of the antiozonant ethylenediurea(EDU0,EDU400)and exposed them to ambient O_(3)(AOZ;08:00-18:00≈30 nmol mol^(-1)) or elevated O_(3)(EOZ;08:00-18:00≈60 nmol mol^(-1))over three growing seasons.We found that EDU400 protected saplings against most effects of EOZ,which included extensive visible foliar injury,premature senescence,decreased photosynthetic pigment contents and altered balance between pigments,suppressed gas exchange and biomass production,and impaired leaf litter decay.While EOZ had limited effects on plant growth(suppressed stem diameter),it decreased the total number of buds per plant,an effect that was not observed in the first growing season.These results indicate that responses to EOZ might have implications to plant competitiveness,in the long term,as a result of decreased potential for vegetative growth.However,when buds were standardized per unit of branches biomass,EOZ significantly increased the number of buds per unit of biomass,suggesting a potentially increased investment to bud development,in an effort to enhance growth potential and competitiveness in the next growing season.EDU400 minimized most of these effects of EOZ,significantly enhancing plant health under O_(3)-induced stress.The effect of EDU was attributed mainly to a biochemical mode of action.Therefore,hybrid larch,which is superior to its parents,can be significantly improved by EDU under long-term elevated O_(3) exposure,providing a perspective for enhancing afforestation practices.

关键词： Air pollution Ethylenediurea(EDU) Hormesis Plant protection Tropospheric ozone(o_3) Larix gmelinii var.japonica×L.kaempferi

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Regulation of the growth of sprouting roots of black locust seedlings using root barrier panels

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Journal of Forestry Research 2023年第3期34卷 655-664页

作者： Satoshi Kitaoka Shiro Matsunami Yannan Wang Saki Fujita Kobayashi Makoto Toshiyuki Hirata Takayoshi Koike Graduate School of Agriculture Hokkaido UniversitySapporo 060-8589Japan The Robust Greenhouse Project of Hokkaido University Sapporo 060-8589Japan Present Address:Oyo Cooperation Ltd Sumitomo BuildingTokyo 101-8486Japan Jiangxi Academy of Forestry Nanchang 330029People’s Republic of China Forestry and Forest Products Research Institute Tsukuba 305-8687Japan Field Science Center for Northern Biosphere Hokkaido UniversitySapporo 060-0809Japan Research Faculty of Agriculture Hokkaido UniversitySapporo 060-8589Japan

How can we regulate an invasive alien species of high commercial value?Black locust(Robinia pseudoacacia L.)has a unique capacity for seed dispersal and high germination.Field surveys indicate that black locust increases its growing area with sprouting roots and the elongation of horizontal roots at a soil depth of 10 cm.Therefore,a method to regulate the development of horizontal roots could be eff ective in slowing the invasiveness of black locust.In this study,root barrier panels were tested to inhibit the growth of horizontal roots.Since it is labor intensive to observe the growth of roots in the fi eld,it was investigated in a nursery setting.The decrease in secondary fl ush,an increase in yellowed leafl ets,and the height in the seedlings were measured.Installing root barrier panels to a depth of 30 cm eff ectively inhibit the growth of horizontal roots of young black locust.

关键词： Black locust Horizontal roots Nutrients Root barrier panel Sprouting roots

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Elevated CO2 offsets the alteration of foliar chemicals(n-icosane,geranyl acetate,and elixene)induced by elevated O3 in three taxa of O3-tolerant eucalypts

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Journal of Forestry Research 2021年第2期32卷 789-803页

作者： Eka Novriyanti Qiaozhi Mao Evgenios Agathokleous Makoto Watanabe Yasuyuki Hashidoko Takayoshi Koike Research and Development Institute of Forest Fiber Technology FOERDIAMinistry of Environment and ForestryKotak Pos 4/BKN BangkinangKamparRiau 28411Indonesia Division of Environment and Resources Research Research Faculty of AgricultureHokkaido UniversitySapporoHokkaido 060-8589Japan College of Resources and Environment Southwest UniversityChongqing 400700China Institute of Applied Ecology Key Laboratory of Agrometeorology of Jiangsu ProvinceNanjing University of Information Science and TechnologyNanjing 210044JiangsuChina Institute of Agriculture Tokyo University of Agriculture and TechnologyFuchuTokyo 183-8509Japan Division of Applied Biological Sciences Research Faculty of AgricultureHokkaido Univ.SapporoHokkaido 060-8589Japan Shenzhen Graduate School of Environment and Energy Peking UniversityBeijing 518055China Research Center for Eco-Environmental Science Chinese Academy of ScienceBeijing 100085China

Eucalypts are important forest resources in southwestern China,and may be tolerant to elevated ground-level ozone(O3)concentrations that can negatively affect plant growth.High CO2 may offset O3-induced effects by providing excess carbon to produce secondary metabolites or by inducing stomatal closure.Here,the effects of elevated CO2 and O3 on leaf secondary metabolites and other defense chemicals were studied by exposing seedlings of Eucalyptus globulus,E.grandis,and E.camaldulensis×E.deglupta to a factorial combination of two levels of O3(<10 nmol mol^(−1)and 60 nmol mol^(−1))and CO2(ambient:370μmol mol^(−1)and 600μmol mol^(−1))in open-top field chambers.GC-profiles of leaf extracts illustrated the effect of elevated O3 and the countering effect of high CO2 on compounds in leaf epicuticular wax and essential oils,i.e.,n-icosane,geranyl acetate and elixene,compounds known as a first-line defense against insect herbivores.n-Icosane may be involved in tolerance mechanisms of E.grandis and the hybrid,while geranyl acetate and elixene in the tolerance of E.globulus.Elevated O3 and CO2,singly or in combination,affected only leaf physiology but not biomass of various organs.Elevated CO2 impacted several leaf traits,including stomatal conductance,leaf mass per area,carbon,lignin,n-icosane,geranyl acetate and elixene.Limited effects of elevated O3 on leaf physiology(nitrogen,n-icosane,geranyl acetate,elixene)were commonly offset by elevated CO2.We conclude that E.globulus,E.grandis and the hybrid were tolerant to these O3 and CO2 treatments,and n-icosane,geranyl acetate and elixene may be major players in tolerance mechanisms of the tested species.

关键词： Carbon dioxide Epicuticular wax Eucalypts Secondary metabolism Tropospheric ozone

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Ethylenediurea(EDU)spray effects on willows(Salix sachalinensis F.Schmid)grown in ambient or ozone-enriched air:implications for renewable biomass production

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Journal of Forestry Research 2022年第2期33卷 397-422页

作者： Evgenios Agathokleous Mitsutoshi Kitao Cong Shi Noboru Masui Shahenda Abu-ElEla Kyohsuke Hikino Fuyuki Satoh Takayoshi Koike Department of Ecology School of Applied MeteorologyNanjing University of Information Science and Technology(NUIST)Nanjing 210044People’s Republic of China School of Agriculture Hokkaido UniversitySapporoHokkaido 060-8589Japan Hokkaido Research Center Forestry and Forest Products Research Institute(FFPRI)Sapporo 062-8516Japan School of Environmental Science and Engineering Tiangong UniversityTianjin 300387People’s Republic of China Department of Entomology Faculty of ScienceCairo UniversityGiza 12613Egypt TUM School of Life Sciences Technical University of Munich(TUM)Hans-Carl-von-Carlowitz Platz 285354 FreisingGermany Field Science Center for Northern Biosphere Hokkaido UniversitySapporo 060-0809Japan Research Center for Eco-Environmental Science Chinese Academy of ScienceBeijing 100085People’s Republic of China

Ground-level ozone(O_(3)) is a widespread air pollutant causing extensive injuries in plants.However,its effects on perennial energy crops remain poorly under-stood due to technical difficulties in cultivating fast-growing shrubs for biomass production under O_(3) treatment on the field.Here we present the results of a two-year evaluation in the framework of which willow(Salix sachalinensis F.Schmid)shrubs were exposed to ambient(AOZ)or elevated(EOZ)O_(3) in two successive growing seasons(2014,2015)and treated with 0(EDU0)or 400 mg L^(−1)(EDU400)eth-ylenediurea spray in the second growing season.In 2014,EOZ altered the chemical composition of both top young and fallen leaves,and a novel mechanism of decreasing Mg in fallen leaves while highly enriching it in young top leaves was revealed in shrubs exposed to EOZ.In 2015,EDU400 alleviated EOZ-induced decreases in leaf fresh mass to dry mass ratio(FM/DM)and leaf mass per area(LMA).While EDU400 protected against EOZ-induced suppression of the maximum rate at which leaves can fix carbon(A_(max))in O_(3)-asymptomatic leaves,it did not alle-viate EOZ-induced suppression of the maximum rates of carboxylation(VCmax)and electron transport(J_(max))and chlorophylls a,b,and a+b in the same type of leaves.In O_(3)-symptomatic leaves,however,EDU400 alleviated EOZ-induced suppression of chlorophylls a and a+b,indicating different mode of action of EDU between O_(3)-asymptomatic and O_(3)-symptomatic leaves.Extensive herbivory occurred only in AOZ-exposed plants,leading to suppressed biomass production,while EOZ also led to a similar suppression of biomass production(EDU0×EOZ vs.EDU400×EOZ).In 2016,carry-over effects were also evaluated following cropping and transplantation into new ambient plots.Effects of EOZ in the preceding growing seasons extended to the third growing season in the form of suppressed ratoon biomass production,indicating carry-over effect of EOZ.Although EDU400 protected against EOZ-induced suppression of biomass production when applied in 2015,there was no

关键词： Air pollution Ethylenediurea(EDU) Hormesis Plant protection Tropospheric ozone(O_(3))

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Ethylenediurea(EDU) effects on Japanese larch: an one growing season experiment with simulated regenerating communities and a four growing season application to individual saplings

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Journal of Forestry Research 2021年第5期32卷 2047-2057页

作者： Evgenios Agathokleous Mitsutoshi Kitao Xiaona Wang Qiaozhi Mao Hisanori Harayama William J.Manning Takayoshi Koike Key Laboratory of Agrometeorology of Jiangsu Province Institute of Applied Ecology Nanjing University of Information Science and Technology (NUIST) Division of Environment and Resources Research Research Faculty of Agriculture Hokkaido University Hokkaido Research Center Forestry and Forest Products Research Institute (FFPRI) College of Landscape Architecture and Tourism Hebei Agricultural University College of Resources and Environment Southwest University Ecophysiology Laboratory Department of Plant Ecology Forestry and Forest Products Research Institute (FFPRI) Department of Plant Soil and Insect Sciences University of Massachusetts Shenzhen Graduate School of Environment and Energy Peking University Research Center for Eco-Environmental Science Chinese Academy of Science

J apanese larch(La rix kaempferi(Lamb.) Carr.)and its hybrid are economically important coniferous trees widely grown in the Northern Hemisphere.Ground-level ozone(O3) concentrations have increased since the preindustrial era,and research projects showed that Japanese larch is susceptible to elevated O3exposures.Therefore,methodologies are needed to(1) protect Japanese larch against O3damage and(2) conduct biomonitoring of O3in Japanese larch forests and,thus,monitor O3risks to Japanese larch.For the first time,this study evaluates whether the synthetic chemical ethylenediurea(EDU) can protect Japanese larch against O3damage,in two independent experiments.In the first experiment,seedling communities,simulating natural regeneration,were treated with EDU(0,100,200,and 400 mg L-1) and exposed to either ambient or elevated O3in a growing season.In the second experiment,individually-grown saplings were treated with EDU(0,200 and 400 mg L-1) and exposed to ambient O3in two growing seasons and to elevated O3in the succeeding two growing seasons.The two experiments revealed that EDU concentrations of 200-400 mg L-1could protect Japanese larch seedling communities and individual saplings against O3-induced inhibition of growth and productivity.However,EDU concentrations ≤200 mg L-1did offer only partial protection when seedling communities were coping with higher level of O3-induced stress,and only 400 mg EDU L-1fully protected communities under higher stress.Therefore,we conclude that among the concentrations tested the concentration offering maximum protection to Japanese larch plants under high competition and O3-induced stress is that of 400 mg EDU L-1.The results of this study can provide a valuable resource of information for applied forestry in an O3-polluted world.

关键词： Air pollution Antiozonant Ethylenediurea (EDU) Plant protection Tropospheric ozone (O 3 )

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Ozone disrupts the communication between plants and insects in urban and suburban areas:an updated insight on plant volatiles

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Journal of Forestry Research 2021年第4期32卷 1337-1349页

作者： Noboru Masui Evgenios Agathokleous Tomoki Mochizuki Akira Tani Hideyuki Matsuura Takayoshi Koike Graduate School of Agriculture Hokkaido UniversitySapporoJapan Key Laboratory of Agrometeorology of Jiangsu Province Institute of EcologySchool of Applied MeteorologyNanjing University of Information Science&TechnologyNanjing 210044People’s Republic of China School of Food and Nutritional Sciences University of ShizuokaShizuokaJapan Research Faculty of Agriculture Hokkaido UniversitySapporoJapan Research Center for Eco-Environmental Science CASBeijing 100085People’s Republic of China

Plant-insect interactions are basic components of biodiversity conservation.To attain the international Sustainable Development Goals(SDGs),the interactions in urban and in suburban systems should be better understood to maintain the health of green infrastructure.The role of ground-level ozone(0_(3)) as an environmental stress disrupting interaction webs is presented.Ozone mixing ratios in suburbs are usually higher than in the center of cities and may reduce photosynthetic productivity at a relatively higher degree.Consequently,carbon-based defense capacities of plants may be suppressed by elevated 0_(3) more in the suburbs.However,contrary to this expectation,grazing damages by leaf beetles have been severe in some urban centers in comparison with the suburbs.To explain differences in grazing damages between urban areas and suburbs,the disruption of atmospheric communication signals by elevated 0_(3) via changes in plant-regulated biogenic volatile organic compounds and long-chain fatty acids are considered.The ecological roles of plant volatiles and the effects of 0_(3) from both a chemical and a biological perspective are presented.Ozone-disrupted plant volatiles should be considered to explain herbivory phenomena in urban and suburban systems.

关键词： Biological interactions Elevated O_3 Insect grazing Pollination Plant defense mechanisms

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Does the root to shoot ratio show a hormetic response to stress? An ecological and environmental perspective

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Journal of Forestry Research 2019年第5期30卷 1569-1580页

作者： Evgenios Agathokleous Regina G.Belz Mitsutoshi Kitao Takayoshi Koike Edward J.Calabrese Hokkaido Research Center Forestry and Forest Products Research Institute(FFPRI)Forest Research and Management Organization7 HitsujigaokaSapporoHokkaido 062-8516Japan Research Faculty of Agriculture Hokkaido UniversityKita 9 Nishi 9SapporoHokkaido 060-8589Japan Agroecology Unit Hans-Ruthenberg InstituteUniversity of HohenheimGarbenstra遝 1370599 StuttgartGermany Department of Environmental Health Sciences Morrill IN344University of MassachusettsAmherstMA 01003USA

Root/shoot(R/S)ratio is an important index for assessing plant health,and has received increased attention in the last decades as a sensitive indicator of plant stress induced by chemical or physical agents.The R/S ratio has been discussed in the context of ecological theory and its potential importance in ecological succession,where species follow different strategies for above-ground growth for light or below-ground competition for water and nutrients.We present evidence showing the R/S ratio follows a biphasic dose–response relationship under stress,typical of hormesis.The R/S ratio in response to stress has been widely compared among species and ecological succession classes.It is constrained by a variety of factors such as ontogeny.Furthermore,the current literature lacks dose-response studies incorporating the full dose–response continuum,hence limiting scientific understanding and possible valuable application.The data presented provide an important perspective for new-generation studies that can advance current ecological understanding and improve carbon storage estimates by R/S ratio considerations.Hormetic response of the R/S ratio can have an important role in forestry for producing seedlings with desired characteristics to achieve maximum health/productivity and resilience under plantation conditions.

关键词： Biological plasticity Biphasic response Hormesis Plant stress Root/shoot ratio

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Effects of soil nutrient availability and ozone on container-grown Japanese larch seedlings and role of soil microbes

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Journal of Forestry Research 2020年第6期31卷 2295-2311页

作者： Evgenios Agathokleous Mitsutoshi Kitao Masabumi Komatsu Yutaka Tamai Hideyuki Saito Hisanori Harayama Akira Uemura Hiroyuki Tobita Takayoshi Koike Institute of Ecology Key Laboratory of Agrometeorology of Jiangsu ProvinceSchool of Applied MeteorologyNanjing University of Information Science and Technology(NUIST)Nanjing 210044JiangsuPeople’s Republic of China Hokkaido Research Center Forestry and Forest Products Research Institute(FFPRI)Forest Research and Management Organization7 HitsujigaokaSapporoHokkaido 062-8516Japan Department of Mushroom Science and Forest Microbiology Forestry and Forest Products Research Institute(FFPRI)Matsunosato 1Tsukuba 305-8687Japan Research Faculty of Agriculture Hokkaido UniversityKita 9 Nishi 9SapporoHokkaido 060-8589Japan Department of Plant Ecology Forestry and Forest Products Research Institute(FFPRI)Matsunosato 1Tsukuba 305-8687Japan

The interactive eff ects of ozone,soil nutrient availability and root microorganisms on physiological,growth,and productivity traits were studied for the fi rst time for Japanese larch(Larix kaempferi)seedlings grown in containers over a growing season,using a free air ozoneconcentration enrichment exposure system.High nutrient availability altered leaf and root nutrient dynamics and enhanced plant growth;however,it also enhanced seedling susceptibility to damping-off disease compared to low nutrient availability.Negative eff ects of elevated ozone,as compared with ambient ozone,on leaf gas exchange and plant stem form were neither off set nor exacerbated by soil nutrient availability and root colonizers.Such negative eff ects suggest that elevated ozone may have implications for ecological health even when plant vigor is limited by factors other than ozone.Inoculation of roots with ectomycorrhizae had negligible infl uence on the eff ects of either soil nutrient availability or ozone.However,this lack of eff ect may be upon impeded formation of complete mycorrhizal root tips due to factors other than the manipulated variables.B and Na appeared to have an important role in stress responses,so further studies to examine their link with physiological mechanisms as a function of time.This study provides an important perspective for designing forestry practices to enhance seedling health.

关键词： Ectomycorrhizae Fertilizer Oxidative stress Plant-fungi interaction

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