Relationship of antioxidant enzymes to ozone tolerance in branches of mature ponderosa pine (Pinus ponderosa) trees exposed to long-term, low concentration, ozone fumigation and acid precipitation

Seasonal activity of superoxide dismutase (SOD, EC 1.15.1.1). ascorbate peroxidase (APOD, EC 1.11.1.11) and guaiacol-oxidizing enzymes (GPODs, EC 1.11.1.7) was examined in needles of 12- to 15-year-old ponderosa pine (Pinus ponderosa Laws.) trees which received ozone (O₃) and acid precipitation treatment. Individual branches were enclosed in branch exposure chambers delivering either charcoal-filtered (O₃-reduced) air, ambient air, or air with twice ambient (2 x ambient) concentrations of O₃. Acid precipitation treatments were rain of pH 3.0 or 5.1 or no rain. Changes in antioxidant enzyme activity were not a consistent response to O₃ fumigation or acid precipitation, but when observed, they occurred most often in the O₃-sensitive clone and in symptomatic, fumigated branches. In the second year of fumigation. O₃ fleck symptoms appeared on needles of the sensitive clone as early as July and APOD activities were significantly increased by O₃ at all sampling dates. In the tolerant clone, antioxidant enzyme activities were not significantly changed by O₃ in the first season of fumigation (March to December 1990), not even during an episode when ambient O₃ concentrations reached 125 nl 1^?1 (240 nl 1^?1 in 2x ambient chambers). No foliar symptoms were observed on needles of the tolerant clone during this year. However, in the second year of fumigation (1992), O₃ fleck symptoms were observed on the tolerant clone and APOD activities were significantly increased in previous-year needles. The tolerant clone had SOD, APOD, and GPOD activities at least 40% higher than those of the sensitive clone before fumigation and 65, 178, and 119% higher, respectively, during both years of fumigation. The higher constitutive levels of these enzymes may have protected against foliar injury in 1990, however in 1992 we concluded that the stimulations in antioxidant enzyme activities observed in symptomatic branches of both clones were a consequence of O₃ injury. Total (intra- and extracellular) activities of the antioxidant enzymes did not appear to be good indicators of O₃ tolerance. Phenotypically, the O₃-tolerant clone was much more vigorous and in both years of fumigation, gas exchange rates were 30 to 71% higher than in the sensitive clone (P. D. Anderson, unpublished data). The greater vigor of the tolerant clone may allow more carbon allocation to protective and repair processes which include, but are not restricted to, the turnover of antioxidant enzymes and metabolites.     

Influence of acid rain and ozone on soil heavy metals under loblolly pine trees: A field study

A field growth chamber study was conducted to determine the effects of ozone and simulated acid rain (SAR) on soil heavy metals. Loblolly pine (Pinus taeda L.), grown in open-top chambers, was exposed to three concentrations of ozone (charcoal filtered air with 0.026 µL O₃ L^-1, and two non-filtered treatments in which ozone concentrations were 0.074 µL L^-1 and 0.147 µL L^-1, respectively) and two levels of SAR (pH 3.5 and 5.2). Ozone was applied for 12 h d^-1 for 9 months and acid rain deposition was 125 mm event^-1. After 9 months exposure, soil pH, organic matter and DTPA-extractable heavy metals (Cd, Pb, Zn, Mn, Fe, Cu) were determined on soil samples collected from exposed chambers at two depths (0–15 cm and 15–30 cm). Simulated acid rain decreased the original soil pH. The concentrations of Cd, Pb and Mn at SAR pH 3.5 were significantly higher than at SAR pH 5.2. Ozone did not affect Zn, Fe and Cu, but a significant interaction between pH and O₃ on Mn, Pb and Cd was observed. Due to the poor drainage capacity of this soil, leaching of heavy metals was not observed.     

Net CO2 exchange of Pinus taeda shoots exposed to variable ozone levels and rain chemistries in field and laboratory settings

Net CO₂ exchange rates (CERs) were measured in seedlings of two loblotly pine ( Pinus taeda L.) families following 6- or 13-week exposures to ozone (charcoalfiltered or ambient air + O₃) and acid rain treatments (pH 3.3, 4.5 and 5.2). Ozone exposures (14 or 170 nl l⁻¹) were made in open-top chambers, and in continously stirred tank reactors (14, 160 or 320 nl l⁻¹) located in the field and laboratory, respectively. The CERs of whole shoots were measured in an open infrared gas analysis system at 6 levels of photosynthetic photon flux density (0, 33, 60, 410, 800 and 1660 μmol m⁻² s⁻¹). Treatment effects were not consistent between field- and laboratory-exposed seedlings. Ozone-treated field seedlings exhibited statistically significant reductions in light-saturated CER of 12.5 and 25% when measured at 6 and 13 weeks, respectively. Laboratory seedlings exhibited mixed responses to O₃, with one family showing reduced CER only after 6 weeks of O₃ exposure and the other only after 13 weeks (O₃ >160 nl l⁻¹ for both). After 13 weeks of exposure, pH 3.3, and 4.5 rain treatments enhanced light-saturated CER by an average of 52% over that observed in seedlings exposed to the pH 5.2 treatment. Enhanced CERs due to acid rain were of the same magnitude (3–5 μmol CO₂g⁻¹ s⁻¹) as ozone-induced CER reductions. No differences in dark respiration were detected between treatments. Although ozone and acid rain treatments altered seedling CER, the differences were not translated into altered final plant dry weights over the 13-week exposure period.     

Hydraulic redistribution of water from Pinus ponderosa trees to seedlings: evidence for an ectomycorrhizal pathway

While there is strong evidence for hydraulic redistribution (HR) of soil water by trees, it is not known if common mycorrhizal networks (CMN) can facilitate HR from mature trees to seedlings under field conditions. Ponderosa pine (Pinus ponderosa) seedlings were planted into root-excluding 61-microm mesh barrier chambers buried in an old-growth pine forest. After 2 yr, several mature trees were cut and water enriched in D(2)O and acid fuchsin dye was applied to the stumps. Fine roots and mycorrhizal root tips of source trees became heavily dyed, indicating reverse sap flow in root xylem transported water from stems throughout root systems to the root hyphal mantle that interfaces with CMN. Within 3 d, D(2)O was found in mesh-chamber seedling foliage > 1 m from source trees; after 3 wk, eight of 10 mesh-chamber seedling stem samples were significantly enriched above background levels. Average mesh-chamber enrichment was 1.8 x greater than that for two seedlings for which the connections to CMN were broken by trenching before D(2)O application. Even small amounts of water provided to mycorrhizas by HR may maintain hyphal viability and facilitate nutrient uptake under drying conditions, which may provide an advantage to seedlings hydraulically linked by CMN to large trees.     

酸雨胁迫对苦槠幼苗气体交换与叶绿素荧光的影响

常绿阔叶树苦槠(Castanopsis sclerophylla)是亚热带地带性顶极群落的建群种之一, 在区域森林资源保护和可持续利用方面具有重要的地位。在该区域日益严重的酸雨胁迫下, 研究其对于胁迫的生理生态响应具有重要的理论价值和实践意义。该文以苦槠幼苗为研究材料, 研究了酸雨胁迫对苦槠幼苗光合生理的影响。结果表明: (1)短时间内, pH 2.5处理下的幼苗叶绿素相对含量最低, 且与pH 5.6处理下的存在显著性差异(p < 0.05); 经过一段时间处理后, pH 4.0处理下的叶绿素相对含量最高, 表明低浓度的酸雨会促进叶绿素相对含量的增加。(2) 2007年4月, 光合速率(P_n)、PSⅡ最大光化学效率(F_v/F_m)和PSⅡ的潜在活性(F_v/F₀)在不同酸雨浓度处理下基本无变化。随着酸雨处理时间的延长, pH 2.5处理下的P_n、光饱和点、光补偿点和暗呼吸速率有显著降低, 且与pH 5.6处理下的存在显著性差异(p < 0.05)。pH 2.5处理组与pH 5.6处理组之间的F_v/F_m和F_v/F₀差异性逐渐减小。表观量子效率和气孔导度变化规律不明显。综合来说, 酸雨处理前期, 高浓度的酸雨胁迫对苦槠幼苗叶绿素相对含量、光合生理参数有显著影响, 但随着酸雨处理时间的增加, 酸雨胁迫对苦槠幼苗的影响逐渐减小, 表明其对外界不良环境具有一定的抵御能力和适应能力。     

红松和紫椴叶片暗呼吸及其光抑制性在幼、成树间的差异

叶片暗呼吸是森林碳循环的重要组分,深入分析幼、成树的叶片暗呼吸及其光抑制性的差异,对生态系统总生产力(GPP)的准确估算具有重要意义.本研究以长白山阔叶红松林主要树种(红松和紫椴)的幼树和成树为研究对象,分别测算不同光照下叶片暗呼吸与无光暗呼吸,比较叶片暗呼吸及其光抑制性在幼、成树间的差异,结合幼、成树叶片生理生态参数的对比,对幼、成树叶片暗呼吸及其光抑制性差异的原因进行探讨.结果表明: 两个树种幼树叶片光下暗呼吸的值高于成树,在生长季(6—9月),幼树的值比成树高6.8%～39.6%;两个树种幼树叶片暗呼吸光抑制程度低于成树,幼树叶片暗呼吸光抑制性的值比成树低2.5%～14.1%;红松幼、成树间叶片暗呼吸光抑制性的差异总体高于紫椴幼、成树间叶片暗呼吸光抑制性的差异,差值最高可达18.6%;幼树中较高的光下暗呼吸值和较低的光抑制程度可能与最大净光合速率、比叶面积、气孔导度的变化有关,与叶片氮含量的变化无关.     

酸雨胁迫对亚热带典型树种幼苗生长与光合作用的影响

通过对酸雨胁迫下中国亚热带典型针叶树和阔叶树,马尾松(Pinus massoniana Lamb.)、杉木(Cunninghamia lanceolata (Lamb.) Hook.)、甜槠(Castanopsis eyrei (Champ.) Tutch.)、苦槠(Castanopsis sclerophylla (Lindl.) Schott.)幼苗的光合生理研究,试图判断日益严重的酸雨是否会对中国亚热带森林产生影响,以便更好地对已退化的森林进行恢复.试验结果表明:在酸雨胁迫下,甜槠和杉木幼苗的高生长受到的影响不显著,而对马尾松幼苗的高生长有一定的促进作用;在pH4.0时,酸雨胁迫对甜槠、杉木和马尾松幼苗的地径生长有一定的促进;在不同酸雨处理下,苦槠、甜槠和马尾松幼苗的最大净光合速率随着酸雨浓度的增加而下降,说明这3种植物幼苗的光合能力受到影响,特别是阔叶树幼苗,苦槠和甜槠幼苗的最大净光合速率分别下降了30%和25%.     

An open-air system for exposure of young forest trees to sulphur dioxide and ozone

The development, construction and operation of an open-air fumigation system for exposing young forest trees to controlled concentrations of sulphur dioxide and ozone is described. A computer simulation of gas dispersion was used to design an array of pipework sources which minimized spatial variability in exposure concentrations. Five fumigation plots were constructed using the design and were used to fumigate trees during a 7 year study known as the Liphook Forest Fumigation Project. Rates of gas release were controlled by a small computer to follow predetermined patterns of sulphur dioxide concentration and to maintain an elevation above ambient ozone concentration. Effective control of exposure was demonstrated, and examples of experimentally produced concentration frequency distributions are provided. The advantages and shortcomings of the system are discussed with recommendations for future improvements.     

The metabolic rate of roach in relation to body size and temperature

Standard and routine metabolic rates of roach Rutilus rutilus for a wide size and temperature range (3–200 g, 5–23° C) were analysed by automated, computerized intermittent flow respirometry. The mass exponent b ranged from 0·68 to 0·82 for standard metabolism, and from 0·65 to 0·92 for routine metabolism depending on the experimental temperature. For routine metabolism b was lowest at 10° C. At both decreasing and increasing temperatures, b increased significantly. Roach were exponentially temperature-dependent for both metabolic levels. For roach <20 g, however, an asymptotic relationship was observed between temperature and routine metabolic rate. The 'flattening of the curve' in the latter case may be explained by reduced spontaneous activities at the lower threshold of the preferred temperature range.     

模拟酸雨胁迫对马尾松和杉木幼苗土壤呼吸的影响

利用LI-8100测定模拟酸雨不同处理下(pH2.5、4.0和5.6)盆栽马尾松(Pinus Massoniana)和杉木幼苗(Cunninghamia lanceolata)的土壤呼吸速率及土壤温度、含水量,研究酸雨对其土壤呼吸的影响.结果表明:模拟酸雨喷淋下马尾松和杉木土壤pH值呈现下降的趋势且下降幅度同酸雨酸度呈现正相关性;马尾松和杉木各个处理下土壤呼吸速率季节变化显著,且同地下10cm土壤温度季节变化趋势一致,pH2.5处理下的土壤呼吸速率平均值分别为1.79μmol · m-2 · s-1和1.12μmol · m-2 · s-1,比对照组(pH5.6)土壤呼吸速率平均值1.57μmol · m-2 · s-1和1.54μmol · m-2 · s-1分别高14%和低39%;马尾松和杉木各个处理下土壤呼吸速率同10cm土壤温度之间均呈现显著的指数关系(P<0.001),与5cm土壤含水量之间相关性不明确;在P=0.05水平上进行多元回归分析,可以得到土壤呼吸速率同土壤温度和含水量的综合拟合方程,和单因素(温度、含水量)拟合相比能够更好地解释土壤呼吸的变化情况;马尾松和杉木在pH2.5和4.0处理下的土壤呼吸温度系数Q10值分别为1.36、2.01和1.51、2 25,同对照组1.14和1.58相比,均有明显差异,且两者Q10值的变化呈先增大后减小的趋势.这证明酸雨是影响马尾松和杉木土壤CO2通量的一个重要因素.     

Growth and maintenance respiration in leaves of northern red oak seedlings and mature trees after 3 years of ozone exposure

A two-component model of growth and maintenance respiration is used to study the response of northern red oak (Quercus rubra L.) seedlings and 32-year-old trees to sub-ambient (10 μmol h; cumulative dose based on 7 h daily mean), ambient (43 μmol h), and twice-ambient (85 μmolh) ozone. The relative growth rates (RGR) of leaves sampled from seedlings and trees were similar across treatments, as were specific leaf respiration rates (SRR). Growth coefficients estimated from the SRR versus RGR relationship averaged 25-3 mol CO2 kg^?1 leaf dry mass produced for seedlings and 21-5 mol kg^?1 for trees. Maintenance coefficients ranged from 0-89 to 1-07 mol CO₂ kg^?1 leaf dry mass d^?1 for seedlings and from 0-64 to 0-84 mol kg-1 d^?1 for trees. Neither coefficient was affected by ozone. Leaves sampled throughout the growing season also showed little response of respiration to ozone. This occurred despite a 30% reduction in net photosynthesis for trees grown at twice-ambient ozone. These results suggest that growth and maintenance respiration in young northern red oak leaves are not affected by ozone and that in older leaves injury can occur without a parallel increase in so-called ‘maintenance’ respiration.     

On the response of pigments and antioxidants of Pinus halepensis seedlings to Mediterranean climatic factors and long-term ozone exposure

An experiment was carried out in open-top chambers located in eastern Spain. One-yr-old Pinus halepensis Mill. seedlings were exposed during three consecutive summers to the following ozone (O₃) treatments: charcoal-filtered air (CFA), non-filtered air (NFA) or non-filtered air plus 40 nl l⁻¹ O₃, 9 h d⁻¹, 5 d wk⁻¹ (NFA+40). Seasonal variations in Aleppo pine performance were observed since reductions in chlorophyll and cellular peroxidase levels associated with increases in superoxide dismutase activity, were recorded during the summer. Similarly, a reduction in epoxidation state was found at midday during the summer, derived from an activation of the xanthophyll cycle associated to an increment in radiation and temperature levels.
The first O₃-induced effects were recorded in previous-year needles (1991) during the first summer exposure as an increase in extracellular and total peroxidase activities and in zeaxanthin levels in the NFA+40 treatment along with a trend to a higher SOD activity in this treatment. A carry-over effect was detected since a lower winter recovery of chlorophyll levels was found in the NFA+40 seedlings along with a reduction of xanthophyll levels. A reduction in chlorophyll levels was observed in the previous-year needles (1992) from the NFA+40 treatment at the end of the second fumigation period. Realistic ozone exposures induced alterations in plant antioxidative systems and plant pigments as shown in this paper. These observations together with the reductions in stomatal conductance and net photosynthesis recorded in the same experiment, indicate that Aleppo pine is a species sensitive to ozone.     

酸雨和UV-B对玉米幼苗光合速率和抗氧化酶活性的影响

以玉米( Zea mays L.)的两个品种渝糯7号和苏玉糯为研究材料, 研究酸雨与UV-B辐射单一因子和复合处理对植物光合作用及抗氧化酶活性的影响机理, 模拟酸雨[pH6.5(A0), 4.5(A1)和3.5(A2)]和增强UV-B辐射[0(B0), 2.88(B1)和5 76(B2)kJ·m-2·d-1]及其复合作用下, 玉米叶片光合速率, 膜脂过氧化程度, 抗氧化酶活性的变化.结果表明, 与对照相比, 酸雨或增强UV-B辐射单因素处理下两个品种的玉米幼苗所测生理指标有相似的变化, 基本表现为: 净光合速率和过氧化氢酶(CAT)活性降低, 质膜透性与丙二醛(MDA)含量增加, 过氧化物酶(POD)活性亦增加, 超氧化物歧化酶(SOD)活性随处理时间延长先升高后降低.复合处理下, 两个品种的玉米幼苗所测生理指标与对照相比也与单因素处理表现相似, 且随处理组合的不同, 复合胁迫下各指标的效果可能表现为协同效应, 但更多地表现为拮抗作用.以上结果说明, 模拟UV-B辐射升高和酸雨pH值降低促进了玉米叶片活性氧的代谢速率, 导致了抗氧化酶系统的紊乱, 抑制了光合作用.在复合处理下, 两个因子对不同指标在不同处理时间和复合水平下, 表现出一定的协同或拮抗效应可能与作物对环境变化产生一定的抗逆性和适应性有关.在不同时期两玉米品种对紫外辐射与酸雨敏感性不同.与苏玉糯相比, 渝糯7号对外界环境变化的抗性更大.     

Ecophysiological responses of Japanese forest tree species to ozone, simulated acid rain and soil acidification

In this review, I summarized the results obtained from experimental studies on the ecophysiological responses of Japanese forest tree species to O₃, simulated acid rain and soil acidification. Based on the studies conducted in Japan, exposure to ambient levels of O₃ below 100 nl·l⁻¹ (ppb) for several months is sufficient to inhibit dry matter production and net photosynthesis of sensitive Japanese forest tree species such as Siebold's beech and Japanese zelkova. On the other hand, exposure to simulated acid rain with a pH of 4.0 or above for several months cannot induce any adverse effects on dry matter production and physiological functions of Japanese forest tree species. However, when the pH of simulated rain or fog is lowered below 4.0, negative effects appear on dry matter production and physiological functions such as transpiration in several sensitive Japanese forest tree species such as Japanese fir and Nikko fir. Based on limited information, it may be concluded that (1) Al dissolved into soil solution is the most important limiting factor for dry matter production, physiological functions and nutrient status of Japanese forest tree species grown in acidic soil, (2) the (Ca+Mg+K)/Al molar ratio in soil solution is a useful indicator to evaluate and predict the effects of soil acidification due to acid deposition on whole-plant dry matter production of Japanese forest tree species at the present time and in the future, and (3) Japanese coniferous tree species such as Japanese cedar and red pine are relatively sensitive to a reduction in (Ca+Mg+K)/Al molar ratio in soil solution compared with European forest tree species such as Norway spruce.