Longevity and senescence of needles in Pinus cembra L.

Summary Needle development has been investigated in Pinus cembra at several locations in the Grisons. Special attention has been paid to longevity and senescence. Despite large variations from one tree to another, longevity (up to 12 years) appeared to increase with increasing altitude. With increasing age, contents of nutrients such as N, P, K and, to a lesser extent, Mg and S tend to decrease. Photosynthetic activity did not decline with increasing age of short shoots. Senescence begins in mid-August in needles of the oldest age classes. It is associated with the mobilization and withdrawal of large proportions of N, P, K and also Mg and S, but Ca appears to be immobile. The extent of yellowing is irregular in individual trees but needle losses on a large scale are followed by reduced losses in subsequent years.     

Intra-annual tracheid production in balsam fir stems and the effect of meteorological variables

Tracheid production of balsam fir in the Québec boreal forest (Canada) was studied by repeated cell analysis to investigate the influence of meteorological variables during the growing seasons 1998 to 2000. Wood micro-cores were extracted on a weekly basis throughout the growing season and sections were prepared in order to count the total number of cells produced. From the weekly cell number obtained, the rate of tracheid production was calculated and correlated with meteorological variables. The average total number of cells produced per year was reasonably uniform, increasing only from 36.6 in 1998, to 41.1 in 2000. However, different cell production rates were noted during the growing season. Regression analysis revealed that the cell production rate was largely dependent on minimum air and soil temperature during most of the cell production period. Mean and maximum temperature had less influence on cell production. Moreover, the influence of temperature was higher during earlywood production mainly from the end of May to mid-July. Lagging the weather data by 1–5 days decreased the relationship between temperature and cell production, showing the high correspondence with the same interval where cell production was measured. These results suggest a fast response of the cambium to temperature variation during tree-ring formation.     

Hemiparasite abundance in an alpine treeline ecotone increases in response to atmospheric CO2 enrichment

Populations of the annual hemiparasites Melampyrum pratense L. and Melampyrum sylvaticum L. were studied at the treeline in the Swiss Alps after 3 years of in situ CO₂ enrichment. The total density of Melampyrum doubled to an average of 44 individuals per square meter at elevated CO₂ compared to ambient CO₂. In response to elevated CO₂, the height of the more abundant and more evenly distributed M. pratense increased by 20%, the number of seeds per fruit by 21%, and the total seed dry mass per fruit by 27%, but the individual seed size did not change. These results suggest that rising atmospheric CO₂ may stimulate the reproductive output and increase the abundance of Melampyrum in the alpine treeline ecotone. Because hemiparasites can have important effects on community dynamics and ecosystem processes, notably the N cycle, changing Melampyrum abundance may potentially influence the functioning of alpine ecosystems in a future CO₂-rich atmosphere.     

Tracheid production in response to indole-3-acetic acid varies with internode age in Pinus sylvestris stems

Summary Different concentrations of indole-3-acetic acid (IAA) in lanolin were applied to the cambial region of approximately 10- and 34-year-old internodes in the main stem of Pinus sylvestris (L.) trees during the tracheid production period. After 5 weeks of treatment, the radial width of xylem produced in both ages of internode was positively related to exogenous IAA concentration measured at 0, 1 and 3 cm directly below the application site. Tracheid production in response to exogenous IAA in the 34-year-old internode was approximately one-half of that in the 10-year-old internode. The endogenous IAA level in the 7-, 17- and approximately 34-year-old internodes of similar trees was measured by radioimmunoassay, using gas chromatography-selected ion monitoring-mass spectrometry for validation. No consistent relationship was found between xylem radial width and IAA concentration. The data indicate that the cambium's ability to respond to exogenous IAA is qualitatively the same in 1-year-old shoots and older internodes. However, as the internode ages, there is a decrease in the extent of the response and in the optimal IAA level for inducing tracheid production.     

Seasonal and spatial variation of woody tissue respiration in a <Emphasis Type="Italic">Pinus cembra</Emphasis> tree at the alpine timberline in the central Austrian Alps

Total stem, branch, twig, and coarse root respiration (R_t) of an adult Pinus cembra tree at the alpine timberline was measured continuously at ten positions from 7 October 2001 to 21 January 2003 with an automated multiplexing gas exchange system. There was a significant spatial variability in woody tissue respiration when expressed per unit surface area or per unit sapwood volume. Surface area related maintenance (R_m) respiration at 0°C ranged between 0.109 and 0.643 mol m^–2 s^–1 and there was no clear trend with respect to tissue type and diameter. Sapwood volume based R_m at 0°C by contrast, varied between 2.5 mol m^–3 s^–1 in the stem and 193.2 mol m^–3 s^–1 in thin twigs in the upper crown. Estimated Q₁₀ values ranged from 1.7 to 3.1. These Q₁₀ values were used along with R_m at 0°C and annual woody tissue temperature records to predict annual total R_m. Annual total R_m accounted for 73±6% of annual R_t in 2002.     

植物群落类型和围栏封育对高山林线岷江冷杉幼苗成活的影响

林线树种幼苗的生长紧密关系着高山林线的形成。通过室内发芽实验测定了岷江冷杉种子萌发能力。2015年春,在川西巴郎山岷江冷杉林线之上的高山林线交错带,通过播种处理(播种和不播种),研究了不同植物群落类型(灌木和草地群落)和围栏处理(围栏和不围栏)对岷江冷杉幼苗成活率的影响,以期为进一步探讨高山林线形成机制提供科学参考。结果表明:(1)岷江冷杉室内发芽率31.4%,成活率随时间下降。(2)灌木群落的全年日平均空气温湿度显著高于草本群落,而两群落的全年日平均土壤温度无显著差异。(3)未播种样地没有发现岷江冷杉幼苗,说明林线交错带缺乏岷江冷杉种子。(4)与不围栏样地相比,围栏封育分别显著提高了岷江冷杉幼苗成活率2.0%(2015年)和2.2%(2016年);与灌木群落相比,草本群落中岷江冷杉幼苗成活率显著提高0.8%(2015年)和1.2%(2016年),说明灌木对林线交错带幼苗更新具有更强的竞争作用。围栏处理下,草本群落中岷江冷杉幼苗成活率显著高于灌木群落;不围栏处理下,草本群落中岷江冷杉幼苗成活率显著低于灌木群落,2017年在不围栏样地没有幼苗存活,说明动物干扰对林线交错带森林更新有阻碍作用。研究表明,高山林线交错带森林更新,受种子、植物竞争和动物干扰(例如,放牧)等多方面因素的共同影响,可以通过播种、植物剔除(去除竞争)和围栏等促进幼苗更新和成活。     

Daily dynamics in xylem cell radial growth of Scots pine (Pinus sylvestris L.)

Daily dynamics of radial cell expansion during wood formation within the stems of 25-year-old Scots pine trees (Pinus sylvestris L.), growing in field conditions, were studied. The samples of forming wood layers were extracted 4 times per day for 3 days. Possible variations in the growth on different sides of the stem, duration of cell development in radial cell expansion phase and dynamics of cell growth in this phase were taken into account. The perimeters of tracheid cross-sections as a reflection of primary cell wall growth were the criterion of growth in a radial direction. For the evaluation of growing cell perimeters a special system for digital processing and image analysis of tracheid cross-sections of the forming wood was used. Growth rate for certain time intervals was estimated by the change in the relation of the perimeter of each observed cell in each of ten tracheid rows in each of 12 trees to the perimeter of the xylem cell of the same row before the expansion. Temporal differences in average values of the relations were estimated by Analyses of Variance. The existence of daily dynamics of Scots pine xylem cell radial growth has been proved. Intensive growth of pine tracheids has been shown to occur at any time of the day and to depend on the temperature regime of the day and the night as well as water supply of stem tissues. Moreover, reliable differences (P = 0.95) in the increment of cell walls during tracheid radial expansion have been found. Pulsing changes of the water potentials both of the cell and the apoplast, as the reason for the fluctuations of radial cell growth rate, were discussed.     

Some quantitative effects of indoleacetic acid on the wood production and tracheid dimensions of Picea

The diameter and wall thickness of tracheids produced after indoleacetic acid treatment were not significantly different from those of the intact controls, for the first few weeks after treatment of disbudded shoots of Picea abies Karst. and Picea sitchensis (Bong.) Carr. However, lateral application of indoleacetic acid (IAA) to intact shoots increased both tracheid diameter and wall thickness; it is suggested that IAA acted synergistically with another endogenous growth regulator, which was also removed by disbudding. Increase in wall thickness after exogenous IAA was associated with increase in duration of the wall thickening phase of tracheid differentiation; this is discussed in relation to the seasonal change from early to latewood. Cambial dormancy was induced by disbudding during active wood production. Since this occurred with or without the presence of current leaves, it is concluded that in Picea continued cambial activity depends upon supply of auxin from the buds, and cannot be supplied from expanded leaves or from the internode itself. Neither indoleacetic acid nor gibberellic acid stimulated renewed cambial activity when applied after the cessation of wood production. With both disbudded and intact shoots, the effectiveness of exogenous IAA declined with time, probably due to decreasing penetration through callus developing at the wounded surface. It is suggested that this apparent change in IAA effectiveness may explain some discrepancies between the results of previous observers.Abbreviations IAA Indoleacetic acid - GA₃ Gibberellic acid     

Carbon allocation in shoots of alpine treeline conifers in a CO<Subscript>2</Subscript> enriched environment

With a new approach we assessed the relative contribution of stored and current carbon compounds to new shoot growth in alpine treeline conifers. Within a free air CO₂ enrichment experiment at the alpine treeline in Switzerland, ¹³C-depleted fossil CO₂ was used to trace new carbon in the two tree species Larix decidua L. and Pinus uncinata Ramond over two subsequent years. The deciduous L. decidua was found to supply new shoot growth (structural woody part) by 46% from storage. Surprisingly, the evergreen P. uncinata, assumed to use current-year photosynthates, also utilized a considerable fraction of storage (42%) for new wood growth. In contrast, the needles of P. uncinata were built up almost completely from current-year photosynthates. The isotopic composition of different wood carbon fractions revealed a similar relative allocation of current and stored assimilates to various carbon fractions. Elevated CO₂ influenced the composition of woody tissue in a species-specific way, e.g. the water soluble fraction decreased in pine in 2001 but increased in larch in 2002 compared to ambient CO₂. Heavy defoliation applied as an additional treatment factor in the second year of the experiment decreased the lipophilic fraction in current-year wood in both species compared to undefoliated trees. We conclude that storage may play an important role for new shoot growth in these treeline conifers and that altered carbon availability (elevated CO₂, defoliation) results in significant changes in the relative amount of mobile carbon fractions in woody tissue. In particular, stored carbon seems to be of greater importance in the evergreen P. uncinata than has been previously thought.     

Characterization of white zones produced on Pinus radiata wood chips by Ganoderma australe and Ceriporiopsis subvermispora

White zones produced on biodegraded Pinus radiata wood chips were characterized by micro-localized-FTIR (Fourier Transformed Infra Red) spectroscopy and scanning electron microscopy. Both techniques permitted assignment of the white zones to a selective lignin removal process. Although both fungi studied have degraded lignin selectively in these restricted superficial areas, chemical analysis of the wood chips indicated that Ganoderma australe removed 16% of the initial amount of glucan at the 20% weight loss level. Ceriporiopsis subvermispora did not remove glucan at weight loss values below 17%. Prolonged biodegradation resulted in reduction of white zones by G. australe, and increased white zones from C. subvermispora decayed samples. This revised version was published online in July 2006 with corrections to the Cover Date.     

Elevated CO2 concentration and temperature effects on the partitioning of chemical components along juvenile Scots pine stems (Pinus sylvestris L.)

The effects of doubled ambient [CO₂] and different temperature levels on young Pinus sylvestris growing in phytotron chambers were studied. Five chambers were supplied with ~380 (‘ambient air’) and five with ~700 μmol mol⁻¹ CO₂ (‘elevated [CO₂]’). Temperature levels in the chambers ranged in increment steps of 2°C from −4°C to +4°C relative to the long-term monthly (day and night) average air temperature levels in Berlin–Dahlem. Substrate was medium fertile; soil moisture and air humidity were kept constant. After three vegetation periods twigs and stems were harvested, weighed, homogenized, and analyzed chemically. There was no significant temperature effect on wood mass accumulation, clearest positive [CO₂] effect occurred in the youngest twigs. In total, wood mass increased by 28.5% at doubled ambient [CO₂]. N-contents (percentage) decreased at elevated [CO₂] in the uppermost stem sections and not in twig wood causing wider C/N ratios in total. In response to elevated temperature, N-contents decreased slightly in twigs (~0.3%). Traces of free glucose, fructose and sucrose, which decreased from the top to the bottom, were found in stem wood, in contrast to traces of starch that increased from the top to the bottom. In response to elevated [CO₂] only a little more (0.05%) was accumulated in the top shoot and in tendency; glucose, fructose, and sucrose contents were lower at the bottom of stems as compared to the control. There was no obvious response of these non-structural carbohydrates to elevated temperature except for starch that decreased to half of the content from the lowest to the highest temperature level. Among the hemicellulose compounds, rhamnose and arabinose declined from the top shoot to the bottom of stem, whereas 4-O-methyl-d-glucuronic-acid, mannose, and xylose increased. Contents (percentage) of galactose remained approximately stable along the stem. The clearest positive effect of elevated [CO₂] along the whole stem was found for mannose with differences of 0.6–0.3%. In contrast to rhamnose and arabinose that showed a negative response to elevated [CO₂], mannose was reduced towards the uppermost stem sections. The 4-O-methyl-d-glucuronic-acid was slightly lowered at the bottom, and galactose and xylose showed no [CO₂] response. The only hemicellulose compound which reacted to temperature elevation was galactose. It increased slightly (~0.1% per 1°C). Cellulose and lignin (Klason) behaved oppositely: cellulose increased and lignin decreased from the top to the bottom. These structural components behaved reversely also in response to elevated [CO₂]. In stem parts above the bottom section, cellulose content was slightly higher at elevated [CO₂], and lignin content was slightly lower at the bottom. Lignin reacted to temperature elevation by a very slight increase on the average (~0.1% per one 1°C). Cellulose, however, decreased by ~0.2% per 1°C temperature elevation. The importance of persistent sinks of carbon in woody plant parts is discussed in respect to the greenhouse effect.     

Vegetation patterns at the alpine treeline ecotone: the influence of tree cover on abrupt change in species composition of alpine communities

Aims: The upper elevation limit of forest vegetation in mountain ranges (the alpine treeline ecotone) is expected to be highly sensitive to global change. Treeline shifts and/or ecotone afforestation could cause fragmentation and loss of alpine habitat, and are expected to trigger considerable alterations in alpine vegetation. We performed an analysis of vegetation structure at the treeline ecotone to evaluate whether distribution of the tree population determines the spatial pattern of vegetation (species composition and diversity) across the transition from subalpine forest to alpine vegetation. Location: Iberian eastern range of the Pyrenees. Methods: We studied 12 alpine Pinus uncinata treeline ecotones. Rectangular plots ranging from 940 to 1900 m² were placed along the forest‐alpine vegetation transition, from closed forest to the treeless alpine area. To determine community structure and species distribution in the treeline ecotone, species variation along the forest‐alpine vegetation transition was sampled using relevés of 0.5 m² set every 2 m along the length of each plot. Fuzzy C‐means clustering was performed to assess the transitional status of the relevés in terms of species composition. The relation of P. uncinata canopy cover to spatial pattern of vegetation was evaluated using continuous wavelet transform analysis. Results: Vegetation analyses revealed a large degree of uniformity of the subalpine forest between all treeline ecotone areas studied. In contrast, the vegetation mosaic found upslope displayed great variation between sites and was characterized by abrupt changes in plant community across the treeline ecotone. Plant richness and diversity significantly increased across the ecotone, but tree cover and diversity boundaries were not spatially coincident. Conclusions: Our results revealed that no intermediate communities, in terms of species composition, are present in the treeline ecotone. Ecotone vegetation reflected both bedrock type and fine‐scale heterogeneity at ground level, thereby reinforcing the importance of microenvironmental conditions for alpine community composition. Tree cover did not appear to be the principal driver of alpine community changes across the treeline ecotone. Microenvironmental heterogeneity, together with effects of past climatic and land‐use changes on ecotone vegetation, may weaken the expected correlation between species distribution and vegetation structure.     

Effects of environmental factors on wood formation in Scots pine stems

Summary To find the optimal conditions for growth and development of tracheid walls in Scots pine stems the effects of temperature and precipitation on xylem cell production by the cambium, radial cell expansion and secondary wall thickening have been studied. The observations were carried out on 10 specially chosen 50 to 60-year-old trees, growing in central Siberia, over 2 seasons. The data on the number of cells in differentiation zones and mature xylem along radial rows of tracheids, radial and tangential sizes of tracheids and their lumens were used for calculating cambial activity, the rates and durations of cell development in the zones, and both the thickness and cross sectional areas of tracheid walls. The mean day, mean maximal diurnal and mean minimal nocturnal temperatures have been shown by correlation and regression analyses to affect differentially separate stages of cytogenesis. The temperature influenced the initial division the side of xylem and radial cell expansion mainly in May–June, while the influence of precipitation increased in July–August. Throughout all seasons it was the temperature that had the main influence on the biomass accumulation in cell walls. Optimal values of temperature and precipitation for cell production by cambium, radial cell expansion and secondary wall thickening have been calculated. The data are discussed in connection with productivity and quality of wood.     

Effect of a synthetic pyrethroid on the growth of ectomycorrhizal fungi and mycorrhiza formation in Pinus patula

Th effect of fenvalerate 20% EC (a synthetic pyrethroid) on the growth of two ectomycorrhizal fungi, Thelephora terrestris and Laccaria laccata, was studied in vitro. In the case of T. terrestris, mycelial growth was promoted at lower concentrations of the insecticide but inhibited at higher concentrations; mycelial growth of L. laccata decreased with increase in concentration. The effect of the insecticide at three concentrations (0.25, 0.5, 1.5 ppm) was also studied on ectomycorrhizal formation in 4-month-old seedlings of Pinus patula in nursery trials. At lower concentrations the insecticide had no significant effect on mycorrhizal formation.     

The effects of flooding on the formation of ectomycorrhizae in Pinus sylvestris seedlings

Pinus sylvestris seedlings, grown in a vertical petri dish system, were inoculated with five different mycorrhizal fungi. Half of the root system in the petri dish was subjected to periodic flooding, and mycorrhizal colonization was studied. Thelephora terrestris, Laccaria laccata, and Hebeloma crustuliniforme were not sensitive to flooding, whereas Suillus flavidus and S. bovinus were highly sensitive. The latter failed to colonize the root even when flooded for only 2 min per day four times a week.     

Mycorrhizal associations in Pinus massoniana Lamb. and Pinus elliottii Engel. inoculated with Pisolithus tinctorius

Dichotomous mycorrhizas were induced in Pinus massoniana and Pinus elliottii seedlings inoculated with Pisolithus tinctorius growing under non-axenic conditions. Six months after inoculation, Pinus massoniana seedlings exhibited a higher degree of infection, bore more mycorrhizas and had developed more abundant extramatrical mycelium than seedlings of Pinus elliottii. Nevertheless, seedlings of Pinus massoniana were stunted and exhibited chorosis of the needles, indicating a possible nutrient deficiency. Histological examination of these pine mycorrhizas showed an ectomycorrhizal association typical of gymnosperms with an intercellular Harting net penetrating between several layers of cortical cells close to the endodermis. However, strong polyphenolic reactions, intracellular hyphae and wall modifications were occasionally observed, indicating that both host-tissue incompatibility and ectendomycorrhizal association can occur in pine species under stressed conditions.     

The Cambial Activity and on Which the Effects of Exogenous IAA in the Stem of Pinus sylvestris L.

The dormant cambial zone consisted of 5–6 cell layers in the main stem of Pinus sylvestris L. trees that were ca. I00 years old. Time of cambial reactivation was comparable at one (bottom) and 8 (top) meters above the ground. In spring, when the cambium reactivated, the number of cambial cells slightly increased and phloem cells were formed. The production of xylem cells followed 3–4 weeks later. The formation of xylem cells decreased, whereas that of phloem cells increased between late June and early July. Cambial reaction in 1-year-old cuttings that were debudded and treated apically with IAA in lanolin was similar to that in the ca. 100-year-old main stem. However, in debudded cuttings treated with plain lanolin, the number of cells in the carnbial zone decreased during the first week of culture, and only a few phloem cells were formed. Later, the fusiform cambial cells of the cambial zone were divided transversely and lost their typical morphology. It is proposed that some factor(s) from roots may stimulate the initiation of cambial cell division, because when the cambium reactivated, the number of cambial cells slightly increased in the ca. 100-year-old main stem, but decreased in the 1-year-old cuttings.     

Gibberellin-induced formation of tension wood in angiosperm trees

After gibberellin had been applied to the vertical stems of four species of angiosperm trees for approximately 2 months, we observed eccentric radial growth that was due to the enhanced growth rings on the sides of stems to which gibberellin had been applied. Moreover, the application of gibberellin resulted in the formation of wood fibers in which the thickness of inner layers of cell walls was enhanced. These thickened inner layers of cell walls were unlignified or only slightly lignified. In addition, cellulose microfibrils on the innermost surface of these thickened inner layers of cell walls were oriented parallel or nearly parallel to the longitudinal axis of the fibers. Such thickened inner layers of cell walls had features similar to those of gelatinous layers in the wood fibers of tension wood, which are referred to as gelatinous fibers. Our anatomical and histochemical investigations indicate that the application of gibberellin can induce the formation of tension wood on vertical stems of angiosperm trees in the absence of gravitational stimulus.     

Temporal and vertical changes in the humus form profile during a primary succession of Pinus sylvestris

The development of the humus form profile during a primary succession of Pinus sylvestris has been studied along chronosequences on dunes and in blow-outs. Attention was given to vertical variation within the humus form and how this changes during profile development. The mor-type ectorganic profile features marked vertical gradients of several soil attributes, while its constituting horizons show no or only small changes of chemical properties during succession. These changes in particular involve increasing calcium and nitrogen concentrations in the organic matter. After an initial high rate of organic matter accumulation in the successive organic horizons, these rates are strongly reduced, suggesting the attainment of a dynamic equilibrium within the time span of the chronosequences on dunes and blow-outs. Blow-outs differ from dunes in the sense that they have a lower amount of organic matter and a higher F/H ratio. This different ratio likely relates to microclimatic conditions less conducive to decomposition.An attempt is made to explain the vertical trends in terms of processes affecting the characteristics of the organic horizons. Main conclusions are that the development of the ectorganic profile results from a combined effect of decay dynamics, rhizosphere processes and atmospheric deposition, which cannot be unentangled quantitatively with the data available. Furthermore, the distinction between F and H horizons has morphological rather than chemical or ecological relevance, as major vertical changes occur within the F horizon.