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and in both years of fumigation, gas exchange rates were 30 to 71% higher than in the sensitive clone (P. D. Anderson, unpublished data).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.     

Pezizalean mycorrhizas and sporocarps in ponderosa pine (Pinus ponderosa) after prescribed fires in eastern Oregon, USA

Post-fire Pezizales fruit commonly in many forest types after fire. The objectives of this study were to determine which Pezizales appeared as sporocarps after a prescribed fire in the Blue Mountains of eastern Oregon, and whether species of Pezizales formed mycorrhizas on ponderosa pine, whether or not they were detected from sporocarps. Forty-two sporocarp collections in five genera (Anthracobia, Morchella, Peziza, Scutellinia, Tricharina) of post-fire Pezizales produced ten restriction fragment length polymorphism (RFLP) types. We found no root tips colonized by species of post-fire Pezizales fruiting at our site. However, 15% (6/39) of the RFLP types obtained from mycorrhizal roots within 32 soil cores were ascomycetes. Phylogenetic analyses of the 18S nuclear ribosomal DNA gene indicated that four of the six RFLP types clustered with two genera of the Pezizales, Wilcoxina and Geopora. Subsequent analyses indicated that two of these mycobionts were probably Wilcoxina rehmii, one Geopora cooperi, and one Geopora sp. The identities of two types were not successfully determined with PCR-based methods. Results contribute knowledge about the above- and below-ground ascomycete community in a ponderosa pine forest after a low intensity fire.     

Soil respiration response to three years of elevated CO2 and N fertilization in ponderosa pine (Pinus ponderosa Doug. ex Laws.)

We measured growing season soil CO₂ evolution under elevated atmospheric [CO₂] and soil nitrogen (N) additions. Our objectives were to determine treatment effects, quantify seasonal variation, and compare two measurement techniques. Elevated [CO₂] treatments were applied in open-top chambers containing ponderosa pine (Pinus ponderosa L.) seedlings. N applications were made annually in early spring. The experimental design was a replicated factorial combination of CO₂ (ambient, + 175, and +350 L L^–1 CO₂) and N (0, 10, and 20 g m^–2 N as ammonium sulphate). Soils were irrigated to maintain soil moisture at > 25 percent. Soil CO₂ evolution was measured over diurnal periods (20–22 hours) in October 1992, and April, June, and October 1993 and 1994 using a flow-through, infrared gas analyzer measurement system and corresponding pCO₂ measurements were made with gas wells. Significantly higher soil CO₂ evolution was observed in the elevated CO₂ treatments; N effects were not significant. Averaged across all measurement periods, fluxes, were 4.8, 8.0, and 6.5 for ambient + 175 CO₂, and +350 CO₂ respectively).Treatment variation was linearly related to fungal occurrence as observed in minirhizotron tubes. Seasonal variation in soil CO₂ evolution was non-linearly related to soil temperature; i.e., fluxes increased up to approximately soil temperature (10cm soil depth) and decreased dramatically at temperatures > 18°C. These patterns indicate exceeding optimal temperatures for biological activity. The dynamic, flow-through measurement system was weakly correlated (r = 0.57; p < 0.0001; n = 56) with the pCO₂ measurement method.     

Recognizing site adversity and drought-sensitive trees in stands of bristlecone pine (Pinus longaeva)

Bristlecone pine has been called the world’s oldest known living thing. The tree grows in mountainous terrain near timberline in three southwestern states. Because of the species’ longevity, sensitivity to climatic fluctuations, and unique growth form, it has attracted considerable attention from both scientists and recreationists. This paper discusses bristlecone pine in relation to its environment in east-central Nevada and sets forth criteria for recognizing trees that may be old or sensitive to climatic changes.     

Primers designed to amplify a mitochondrial nad1 intron in ponderosa pine, Pinus ponderosa, limber pine, P. flexilis, and Scots pine, P. sylvestris

The b/c intron of the mitochondrial nad1 gene, was sequenced to characterize the indel region of ponderosa pine, Pinus ponderosa. The sequence in ponderosa pine was aligned with the sequence in Scots pine, Pinus sylvestris, to design seven primers that are useful for sequencing and for revealing size variation in amplified fragments in ponderosa pine, Scots pine, and limber pine, Pinus flexilis. These primers reveal variability in all three species, and the pattern of variability within ponderosa pine is described by a preliminary survey. The indel region of ponderosa pine contains three distinct elements with lengths of 31, 32, and 34 bp. Received: 1 March 2000 / Accepted: 14 April 2000<@head-com-p1a.lf>Communicated by P.M.A. Tigerstedt     

Shoot multiplication from mature trees of Douglas-fir (Pseudotsuga menziesii) and sugar pine (Pinus lambertiana)

Opening of apical and axillary buds of mature Douglas-fir and sugar pine trees was obtained on a newly formulated basal medium (DCR) without growth regulators. Elongation of buds was observed on 1/2 strength DCR with 0.3% activated charcoal (DCR-1). In sugar pine, multiple shoots were obtained when explants on DCR with 0.5 mg/1 BAP for 5–6 weeks were transferred to DCR-1 medium. On subculture, axillary buds again developed when shoots were cultured on DCR with 0.2 mg/1 BAP for Douglas-fir and 0.5 mg/1 BAP for sugar pine. These buds were again elongated on DCR-1 medium. By subculturing 7–10 shoots of Douglas-fir and 2–3 shoots of sugar pine, over 100 shoots can be obtained in a year.Abbreviations BAP N⁶-benzylaminopurine - KN kinetin - NAA -naphthalene acetic acid - IAA Indole-3-acetic acid - MS Murashige-Skoog medium - WPM woody-plant medium     

Palaeoclimate potential of New Zealand Manoao colensoi (silver pine) tree rings using Blue-Intensity (BI)

This study investigates if Blue Intensity (BI) parameters are capable of capturing enhanced climatic signals from a key New Zealand dendrochronological species when compared to ring-width (RW) measurements. Three BI parameters (earlywood mean, latewood mean and maximum latewood) recorded generally superior correlations to temperature than conventional RW measurements and these correlations appeared more stable over time. For all BI parameters, austral summer (Dec-Feb) was identified as the strongest reconstructive window. However, all the BI parameters required greater sample replication than RW to reach widely accepted thresholds of chronology reliability, highlighting the need for further examination into the physiological controls on the BI signal. Despite the need for further research, this pilot study is strongly encouraging for the wider application of BI in Southern Hemisphere dendrochronology.     

Cambial history of Scots pine trees (Pinus Sylvestris) prior and after the Chernobyl accident as encoded in the xylem

Studies were carried out on wood samples collected in October 1997 from breast height of Scots pine trees (Pinus sylvestris) from site located 5 km south from the Chernobyl nuclear power plant. The radioactive contamination at the site was 3.7x10(5) kBq m(-2). These samples of secondary wood were used as an archive of information about the dynamics of a meristematic tissue cambium affected by ionising radiation from the Chernobyl reactor accident. The results show that frequency of the cambial cells events like anticlinal divisions, intrusive growth and cells elimination, was after the Chernobyl accident, about three times higher in comparison to preceding years. The most interesting finding was that after irradiation the length of tracheids increased. This increase is interpreted as an effect of intracambial competition among cells in the initial layer.     

No unintended impacts of transgenic pine (Pinus radiata) trees on above ground invertebrate communities

As part of an investigation into the potential unintended ecological impacts of transgenic trees, invertebrates were sampled from a field trial of transgenic Pinus radiata D. Don carrying the expressed antibiotic resistance marker gene neomycin phosphotransferase II (nptII) along with other genes known to affect reproductive development in plants and from nontransformed control trees. Invertebrate species abundance, richness, diversity, and composition were measured and compared among trees of five different transclones and nontransformed isogenic control trees. Invertebrates were sampled at six-monthly intervals over a period of 2 yr. In total, 19,162 individuals were collected comprising 279 invertebrate recognizable taxonomic units. Total invertebrate populations as well as populations of herbivorous lepidopteran larvae and Hemiptera were compared among transgenic and control trees. The results show that the transclones had no significant unintended influence on species abundance, richness, diversity, or composition for all populations investigated.     

Tree rings reveal extent of exposure to ionizing radiation in Scots pine Pinus sylvestris

Tree growth has been hypothesized to provide a reliable indicator of the state of the external environment. Elevated levels of background ionizing radiation may impair growth trajectories of trees by reducing the annual growth. Such effects of radiation may depend on the individual phenotype and interact with other environmental factors such as temperature and drought. We used standardized growth rates of 105 Scots pine Pinus sylvestris located near Chernobyl, Ukraine, varying in the level of background radiation by almost a factor 700. Mean growth rate was severely depressed and more variable in 1987–1989 and several other subsequent years, following the nuclear accident in April 1986 compared to the situation before 1986. The higher frequency of years with poor growth after 1986 was not caused by elevated temperature, drought or their interactions with background radiation. Elevated temperatures suppressed individual growth rates in particular years. Finally, the negative effects of radioactive contaminants were particularly pronounced in smaller trees. These findings suggest that radiation has suppressed growth rates of pines in Chernobyl, and that radiation interacts with other environmental factors and phenotypic traits of plants to influence their growth trajectories in complex ways.     

Family 34 glycosyltransferase (GT34) genes and proteins in Pinus radiata (radiata pine) and Pinus taeda (loblolly pine)

Using a functional genomics approach, four candidate genes (PtGT34A, PtGT34B, PtGT34C and PtGT34D) were identified in Pinus taeda. These genes encode CAZy family GT34 glycosyltransferases that are involved in the synthesis of cell‐wall xyloglucans and heteromannans. The full‐length coding sequences of three orthologs (PrGT34A, B and C) were isolated from a xylem‐specific cDNA library from the closely related Pinus radiata. PrGT34B is the ortholog of XXT1 and XXT2, the two main xyloglucan (1→6)‐α‐xylosyltransferases in Arabidopsis thaliana. PrGT34C is the ortholog of XXT5 in A. thaliana, which is also involved in the xylosylation of xyloglucans. PrGT34A is an ortholog of a galactosyltransferase from fenugreek (Trigonella foenum‐graecum) that is involved in galactomannan synthesis. Truncated coding sequences of the genes were cloned into plasmid vectors and expressed in a Sf9 insect cell‐culture system. The heterologous proteins were purified, and in vitro assays showed that, when incubated with UDP‐xylose and cellotetraose, cellopentaose or cellohexaose, PrGT34B showed xylosyltransferase activity, and, when incubated with UDP‐galactose and the same cello‐oligosaccharides, PrGT34B showed some galactosyltransferase activity. The ratio of xylosyltransferase to galactosyltransferase activity was 434:1. Hydrolysis of the galactosyltransferase reaction products using galactosidases showed the linkages formed were α‐linkages. Analysis of the products of PrGT34B by MALDI‐TOF MS showed that up to three xylosyl residues were transferred from UDP‐xylose to cellohexaose. The heterologous proteins PrGT34A and PrGT34C showed no detectable enzymatic activity.     

Light rings in Chinese pine (Pinus tabulaeformis) in semiarid areas of north China and their palaeo-climatological potential

Light rings in conifer trees are characterized by a light-coloured, narrow latewood band of thin-walled tracheids. Most reports on light rings have been for subarctic and subalpine regions, and little is known about their occurrence in semiarid areas. Dendrochronological methods were used to date the occurrence of light rings in Chinese pine (Pinus tabulaeformis) in the semiarid region of north China. The anatomical and chemical characteristics and the potential environmental controls of their formation were investigated. Light rings in Chinese pine were dated to the year of their formation. The wall thickness and lumen diameter of the wood cells of light rings and reference rings were distinctly different. However, the configuration of the light-ring latewood cell walls was normal, although they were thinner than average, and their lignification had been completed normally. The climate characteristics that result in light-ring formation appear to be ongoing severe drought from the previous autumn to July of the current year in conjunction with a warm summer, suggesting that light rings can be used as indicators for past drought events.     

A functional-structural model for radiata pine (Pinus radiata) focusing on tree architecture and wood quality

Backgrounds and Aims

Functional–structural models are interesting tools to relate environmental and management conditions with forest growth. Their three-dimensional images can reveal important characteristics of wood used for industrial products. Like virtual laboratories, they can be used to evaluate relationships among species, sites and management, and to support silvicultural design and decision processes. Our aim was to develop a functional–structural model for radiata pine (Pinus radiata) given its economic importance in many countries.

Methods

The plant model uses the L-system language. The structure of the model is based on operational units, which obey particular rules, and execute photosynthesis, respiration and morphogenesis, according to their particular characteristics. Plant allometry is adhered to so that harmonic growth and plant development are achieved. Environmental signals for morphogenesis are used. Dynamic turnover guides the normal evolution of the tree. Monthly steps allow for detailed information of wood characteristics. The model is independent of traditional forest inventory relationships and is conceived as a mechanistic model. For model parameterization, three databases which generated new information relating to P. radiata were analysed and incorporated.

Key Results

Simulations under different and contrasting environmental and management conditions were run and statistically tested. The model was validated against forest inventory data for the same sites and times and against true crown architectural data. The performance of the model for 6-year-old trees was encouraging. Total height, diameter and lengths of growth units were adequately estimated. Branch diameters were slightly overestimated. Wood density values were not satisfactory, but the cyclical pattern and increase of growth rings were reasonably well modelled.

Conclusions

The model was able to reproduce the development and growth of the species based on mechanistic formulations. It may be valuable in assessing stand behaviour under different environmental and management conditions, assisting in decision-making with regard to management, and as a research tool to formulate hypothesis regarding forest tree growth and development.     

长白山阔叶红松林树木短期死亡动态

树木死亡是森林群落动态的重要过程,是多种因素共同作用的结果.本文基于长白山阔叶红松林25 ha样地2004年和2009年两次调查数据,从物种组成、数量特征、径级结构和空间分布等方面分析了5年间样地中死亡树木的特征.结果表明:5年间样地DBH≥1 cm的独立个体树种数由52变为51,3个树种因权有的1个个体死亡而消失,新增2个树种;独立个体数从36,908变为34,926,死亡个体数为4,030,死亡个体数占2004年个体总数的10.9%,新增个体数为2,048,独立个体数净减少1,982;死亡量大的树种其新增量也较大,灌木树种的死亡量和新增量均多于乔木树种;有5个树种的平均胸径减小,44个树种平均胸径增加;从死亡个体的径级结构来看,小径级个体死亡较多,大径级个体死亡少,5 cm以下的死亡个体占总死亡量的81.5%,不同林层的优势树种死亡个体的径级分布与2004年该树种的径级分布基本相同;不同树种死亡个体的空间分布具有较大差异,不同林层的优势树种死亡个体空间格局主要以聚集分布为主,小径级死亡个体在小尺度呈聚集分布,在其他尺度呈随机分布,中径级和大径级死亡个体在各尺度上都呈不同程度的随机分布.     

Paternal cytoplasmic transmission in Chinese pine (Pinus tabulaeformis)

Summary. In this paper, the stages of normal sexual reproduction between pollen tube penetration of the archegonium and early embryo formation in Pinus tabulaeformis are described, emphasizing the transmission of parental cytoplasm, especially the DNA-containing organelles – plastids and mitochondria. The pollen tube growing in the nucellus contained an irregular tube nucleus followed by a pair of sperm cells. The tube cytoplasm contained abundant organelles, including starch-containing plastids and mitochondria. The two sperm cells differed in their volume of cytoplasm. The leading sperm, with more cytoplasm, contained abundant plastids and mitochondria, while the trailing one, with a thin layer of cytoplasm, had very few organelles. The mature egg cell contained a great number of mitochondria, whereas it lacked normal plastids. At fertilization, the pollen tube penetrated into the egg cell at the micropylar end and released all of its contents, including the two sperms. One of the sperm nuclei fused with the egg nucleus, whereas the other one was retained by the receptive vacuole. Very few plastids and mitochondria of male origin were observed around the fusing sperm and egg nuclei, while the retained sperm nucleus was surrounded by a large amount of male cytoplasm. The discharged tube cytoplasm occupied a large micropylar area in the egg cell. In the free nuclear proembryo, organelles of maternal and paternal origins intermingled in the neocytoplasm around the free nuclei. Most of the mitochondria had the same features as those of the egg cell, but some appeared to be from sperm cells and tube cytoplasm. Plastids were obviously of male origin, with an appearance similar to those of the sperm or tube cells. After cellularization of the proembryo, maternal mitochondria became more abundant than the paternal ones and the plastids enlarged and began to accumulate starch. The results reveal the cytological mechanism for paternal inheritance of plastids and biparental inheritance of mitochondria in Chinese pine. Correspondence and reprints: State Key Laboratory of Plant Physiology and Biochemistry, College of Biological Science, China Agricultural University, Beijing 100094, People’s Republic of China.     

Delayed seed dispersal in Pinus torreyana (Torrey pine)

Summary Torrey pine exhibits a pattern of seed release intermediate between an open and closed-cone species. It was found that the cones open at maturity, but that seed fall from some cones continues for up to 13 years. Approximately 77% of the total seed crop was present in age classes one or more years past the time of seed maturity.Serotiny is thought to evolve in response to fire, and partial serotiny at the population level has been attributed to circumstances in which the strong selective pressure of fire is relaxed. We hypothesize that delayed seed dispersal in Torrey pine is a stable trait which is the result of a pattern of large relatively rare crown fires separated by intervals longer than one generation. The reduced serotiny observed in other conifers on headlands and islands in California seems consistent with our hypothesis.     

Tree rings of Scots pine (Pinus sylvestris L.) as a source of information about past climate in northern Poland

Scots pine (Pinus sylvestris) is a very common tree in Polish forests, and therefore was widely used as timber. A relatively large amount of available wood allowed a long-term chronology to be built up and used as a source of information about past climate. The analysis of reconstructed indexed values of mean temperature in 51-year moving intervals allowed the recognition of the coldest periods in the years 1207–1346, 1383–1425, 1455–1482, 1533–1574, 1627–1646, and 1694–1785. The analysis of extreme wide and narrow rings forms a complementary method of examining climatic data within tree rings. The tree ring widths, early wood and late wood widths of 16 samples were assessed during the period 1581–1676. The most apparent effect is noted in the dry summer of 1616. According to previous research and our findings, temperature from February to March seems to be one of the most stable climatic factors which influenced pine growth in Poland. Correlation coefficients in the calibration and validation procedure gave promising results for temperature reconstruction from the pine chronology.     

Changing growth response to wildfire in old‐growth ponderosa pine trees in montane forests of north central Idaho

North American fire‐adapted forests are experiencing changes in fire frequency and climate. These novel conditions may alter postwildfire responses of fire‐adapted trees that survive fires, a topic that has received little attention. Historical, frequent, low‐intensity wildfire in many fire‐adapted forests is generally thought to have a positive effect on the growth and vigor of trees that survive fires. Whether such positive effects can persist under current and future climate conditions is not known. Here, we evaluate long‐term responses to recurrent 20th‐century fires in ponderosa pine, a fire‐adapted tree species, in unlogged forests in north central Idaho. We also examine short‐term responses to individual 20th‐century fires and evaluate whether these responses have changed over time and whether potential variability relates to climate variables and time since last fire. Growth responses were assessed by comparing tree‐ring measurements from trees in stands burned repeatedly during the 20th century at roughly the historical fire frequency with trees in paired control stands that had not burned for at least 70 years. Contrary to expectations, only one site showed significant increases in long‐term growth responses in burned stands compared with control stands. Short‐term responses showed a trend of increasing negative effects of wildfire (reduced diameter growth in the burned stand compared with the control stand) in recent years that had drier winters and springs. There was no effect of time since the previous fire on growth responses to fire. The possible relationships of novel climate conditions with negative tree growth responses in trees that survive fire are discussed. A trend of negative growth responses to wildfire in old‐growth forests could have important ramifications for forest productivity and carbon balance under future climate scenarios.