Long term effect of liming and fertilization on ectomycorrhizal colonization and tree growth in old Scots pine (Pinus sylvestris L.) stands

In this study, we surveyed the long term effects of liming and fertilizing in old Scots pine stands on the ectomycorrhiza (ECM) colonization, tree growth and needle nutrient concentration 35 years later. Four mature stands of Scots pine on low productive mineral soil were limed in 1959 and 1964 with total doses of limestone ranging from 3 to 15 Mg ha^?1 and fertilized with nitrogen (N) in 1970. Thirty-five years after the first liming treatment, all stands were analysed for tree growth and needle nutrient concentrations and two of the stands were also analysed for ECM colonization. ECM colonization increased significantly with liming from 61.5% in the control plots to 88% in the plot with the highest limestone dose. ECM colonization increased with increasing pH in the humus layer from 62% colonization at pH?=?3.5 to 90% at pH?=?6.5 and decreased with increasing amount of extractable phosphorus (P) in the humus. Liming did not affect the frequencies of different ECM morphotypes or dead short root tips, the fine root biomass or necromass. ECM colonization was uncorrelated with needle nutrient concentrations or tree increment. Liming did not significantly affect tree growth. However, nutrient concentrations of current-year needles were affected by prior liming. Ca concentrations in current-year needles increased from approximately 15 mg g^?1 in control treatments to more than 30 mg g^?1 in limed plots, whereas concentrations of Mn, Al, Fe, and in two stands, B, decreased due to liming. In conclusion, liming with doses up to 15 Mg ha^?1 was detectable in stands 35 years after treatment. The liming significantly increased the ECM colonization of Scots pine fine roots, increased the needle nutrient concentration of Ca and decreased the needle concentrations of Mn, Al, and Fe.     

Root biomass changes after drainage and fertilization of a low-shrub pine bog

The stump and root systems of Scots pine (Pinus sylvestris) and field-layer vegetation were sampled before (1984) and three growing seasons after drainage and fertilization (1987) of a low-shrub pine bog. Average below-ground biomass of the field layer was 548 gDW m^–2 in 1984, with no significant treatment effects during experimentation. The stump-plus-root biomass of the pine stands was 1464 gDW m^–2 in the virgin state, and had increased to 1854 gDW m^–2 three years after the NPK-fertilizer treatment. The distribution over fractions also changed with this treatment. The fraction of fine roots ( < 1 mm) in stump-root biomass increased from 4% (56 gDW m^–2) to 11% (196 gDW m^–2), while the other compartments changed less. Total pine root length was 729 mm^–2 in 1984. Root length increased by 94% to 1380 mm^–2 on NPK-fertilized plots. Most of the fine pine roots were in the surface layer (0–10 cm), 79% in 1984 and 88% in 1987, and few pine roots were deeper than 20 cm. Maximum root length of fine pine roots ( < 1 mm) was estimated to be 2710 mm^–2 at about 800 gDW m^–2 (NPK treatment), and the corresponding maximum for small pine roots (=1–10 mm) was 227 mm^–2 at 809 gDW m^–2. Drainage stimulated net growth of fine roots, but this treatment also caused higher mortality rates of small roots. The fine roots responded to fertilization with higher net growth rate, and secondary growth of the large roots ( > 10 mm) was improved. The observed changes in root biomass and structure are explained as strategic adaptations to altered hydrological and nutritional circumstances in the root zone after drainage and fertilization.     

Defoliation effects on enzyme activities of the ectomycorrhizal fungus <Emphasis Type="Italic">Suillus granulatus</Emphasis> in a <Emphasis Type="Italic">Pinus contorta</Emphasis> (lodgepole pine) stand in Yellowstone National Park

Ectomycorrhizal (EM) basidiomycete fungi are obligate mutualists of pines and hardwoods that receive fixed C from the host tree. Though they often share most recent common ancestors with wood-rotting fungi, it is unclear to what extent EM fungi retain the ability to express enzymes that break down woody substrates. In this study, we tested the hypothesis that the dominant EM fungus in a pure pine system retains the ability to produce enzymes that break down woody substrates in a natural setting, and that this ability is inducible by reduction of host photosynthetic potential via partial defoliation. To achieve this, pines in replicate blocks were defoliated 50% by needle removal, and enzyme activities were measured in individual EM root tips that had been treated with antibiotics to prevent possible bacterial activity. Results indicate that the dominant EM fungal species (Suillus granulatus) expressed all enzymes tested (endocellulase D-glucosidase, laccase, manganese peroxidase, lignin peroxidase, phosphatase and protease), and that activities of these enzymes increased significantly (P < 0.001) in response to defoliation. Thus, this EM fungus (one of the more specialized mutualists of pine) has the potential to play a significant role in C, N and P cycling in this forested ecosystem. Therefore, many above-ground factors that reduce photosynthetic potential or divert fixed C from roots may have wide-reaching ecosystem effects.     

Development of needle retention in Scots pine (Pinus sylvestris) in 1957–1991 in northern and southern Finland

The needle trace method was used to study retrospectively the long-term latitudinal variation in needle retention in Scots pine (Pinus sylvestris L.) in Finland. The mean annual summer needle retention (ANR) along the main stem varied from 3.4 to 6.0 needle sets during the period 1957–1991. The lowest values were observed in southern and the highest in northern Finland. The length of the growing season, expressed as the thermal sum (threshold value +5 °C), was negatively correlated with the mean ANR (r=-0.96). The geographical needle retention pattern (NRP) for the period 1957–1991 showed a clearly increasing trend from 1957 to 1969 (southern Finland) and to 1975 (northern Finland); thereafter, the NRP tended to decrease close to its minimum value recorded in 1991. The general level of the NRP was approximately 5.0 needle sets in northern Finland and 3.5–4.0 needle sets in southern Finland. The NRP, with its 6–12 year cycle for southern Finland, was clearly periodical. Differences in the NRP among the ten stands in southern Finland were small, whereas the said periodicity was missing and the differences were high among the stands in northern Finland. The results indicate that variation in the number of needle sets, viz. defoliation of pines, is a normal phenomenon. The role of net carbon assimilation as a regulator of the number of needle sets is discussed.     

Advances in the positional cloning of nodulation genes in soybean

The effect of liming on the decomposition of Norway spruce needle litter was studied in 40–60-year-old Norway spruce stands. Finely-ground limestone had been spread about 30 years ago at a dose of 2 t ha^–1 and reliming was carried out about 20 yr later at a dose of 4 t ha^–1. Needle litter was collected from both control and limed plots, and it was placed in litter bags in the middle of the humus layer of the plot from which they originated, and similarly to the other plot in May. Litter bags were sampled after 4, 12 and 16 months. The site of origin of the needle litter, whether from control plot or from limed plot, affected mainly the early stages of decomposition. Initially the effect of liming was seen as decreased concentration of water soluble material and then, during decomposition, as decreased mass loss and decreased degradation of lignin, and increased C/N ratio. The incubation site, whether the control or the limed plot, did not affect decomposition significantly.Decomposition of Scots pine needles in a young Scots pine plantation was also studied. The treatments were: 2 t ha^–1 of finely-ground limestone and 2.5 t ha^–1 of bark ash spread 8 months before this study. The treatments did not affect decomposition much, but some stimulation of the treatments on decomposition was observed. Compared to spruce needles, the C/N ratio of pine seedles was lower, they contained less lignin and more water soluble material, and decomposed faster in the first summer.     

Defoliation and mycorrhizal symbiosis: a functional balance between carbon sources and below-ground sinks

Herbivory is generally assumed to negatively influence mycorrhizal fungi because of reduced photosynthate to support mycorrhizae following defoliation. We examined effects of 60% and 100% defoliation (excluding current year needles) on tree growth and ectomycorrhizal associations of 10–15 year old Scots pines ( Pinus sylvestris ). Over 98% of short roots were colonized by mycorrhizal fungi, and contrary to expectation, defoliation did not decrease the proportion of living fungi in fine roots. Furthermore, defoliation did not alter the ratios of produced needle biomass to the biomass of fine roots or living fungi in fine roots. The composition of mycorrhizal morphotypes was changed, however, which suggests competition among different mycorrhizal growth forms owing to their carbon demands. We propose that these outcomes are a consequence of a functional balance between carbon sources in plant foliage and below-ground sinks, i.e. growing roots and mycorrhizal associates.     

Feast and famine: previous defoliation limiting survival of pine processionary caterpillar Thaumetopoea pityocampa in Scots pine Pinus sylvestris

This study analyses the consequences of previous defoliation on the survival of the larvae of the pine processionary moth Thaumetopoea pityocampa (Denis and Schiffermüller) feeding on relict Scots pine Pinus sylvestris (L.) ssp. nevadensis Christ in the Sierra Nevada mountains (SE Spain). Egg batches of the pine processionary moth were placed on four groups of Scots pines that underwent different periods of herbivory. The larval survival was related to the nitrogen content, fibre, phenolics and terpenes in the needles. Larval survival was higher in undefoliated pines, lower in pines defoliated two consecutive years, and intermediate in pines defoliated only one year, suggesting a direct relationship between previous defoliation and larval survival. In contrast, none of the characteristics of the needles showed a clear relationship with larval survival. The resulting reduction in larval number also affects the capacity of the larvae to develop during winter, because it hampered nest warming. Thus, previous defoliation limits, although it does not impede, the possibility of repeated defoliation on Scots pine.     

The Complementarity of Extractable and Ester-Bound Lipids in a Soil Profile Under Pine

Extractable and solvent insoluble, ester-bound lipids were analysed in an acid, sandy soil profile under Corsican pine. The n-alkanes and alkanoic acids from the soil profile showed rather poor correlations with those from the pine needles and roots, while the n-alkanol composition in the mineral horizons strongly indicated the presence of lipids derived from a previous grass vegetation. Although the ester-bound lipids (ω-hydroxyalkanoic acids and α,ω-alkanedioic acids (>C₂₄)) suggested that plant sources other than pines were present in the mineral soil horizons their composition was less contaminated and a clear distinction between needle and root input could be discerned. The divergent clustering of soil horizons and plant materials by individual and combined compound classes emphasized the usefulness of both extractable lipids and cutin/suberin in unravelling (past) vegetation and tissue history and contributions to soil organic matter.     

Comparison of Fine Root Dynamics in Scots Pine and Pedunculate Oak in Sandy Soil

In this study, we investigated the relationship between the seasonality of vegetation cover and that of fine root processes in a man-made forest in northern Belgium. Due to their contrasting foliar development, we expected different seasonal patterns of fine root growth and standing biomass between Pedunculate oak (Quercus robur L.), and Scots pine (Pinus sylvestris L.). Biomass and necromass of fine and small roots were estimated by repeated core sampling in February, April, June, August and October 2003. Measurements showed that Pedunculate oaks maintained more live fine roots in winter than Scots pines. However, Scots pines produced more than twice as much fine roots in spring, such that in summer both species had similar root mass. Scots pine root production started before-, but declined during leaf unfolding. Pedunculate oak roots, in contrast, started elongating only after bud break. For both species, fine root production peaked in JuneJuly, but was more than offset by drought-induced mortality at the end of July and early August. Summer drought in 2003 was exceptionally long and intense, significantly reducing leaf area, killing most new roots, and inhibiting root decomposition, such that the obtained results cannot be typical for this forest.     

Effects of defoliation and symbiosis on polyamine levels in pine and birch

 We report the effect of ectomycorrhizal fungi (Suillus variegatus, Paxillus involutus) and defoliation on polyamine concentrations in pine (Pinus silvestris) and birch (Betula pendula) foliage and roots. Symbiotic root tips showed consistently higher concentrations of putrescine than non-symbiotic roots. Partial defoliation had no effect on the polyamine levels in mycorrhizal pine or birch roots. The foliage of mycorrhizal pine seedlings had lower putrescine concentrations and higher spermidine than foliage of non-mycorrhizal plants, and defoliation reversed this pattern. The response to partial defoliation differed in birch foliage: mycorrhizal status had no effect and all new growth after defoliation had higher spermidine levels than in non-defoliated birch. The potential role of polyamines in mycorrhizal symbiosis is discussed. Accepted: 26 February 1997     

松材线虫(Bursaphelenchus xylophilus)入侵对马尾松(Pinus massoniana)林分生长的影响及相关生长模型

松材线虫（Bursaphelenchus xylophilus）是一种松树上发生严重的有害生物,它不仅改变了生态系统的结构和功能,而且改变了系统内生物的原有特性和地理分布。松材线虫及其引起的松树萎蔫病已对中国马尾松林（Pinus massoniana）的树木成长产生了巨大影响。基于此,使用＂每木调查法＂和＂样方法＂,对松材线虫入侵后的马尾松林内松树的各项生长指标因子进行了调查分析,其结果表明：自松材线虫1996年入侵所调查地区的松林后,对于受害松树不管是伐倒木（被伐倒）还是倒木（自然倒地）,其对周围马尾松胸径生长的影响是显著的,而对树高生长的影响不显著。最后建立了一系列的灰色和灰色-马尔可夫链数学模型,其预测结果精度高,可用于今后受害和未受害区马尾松林分因子的生长预测。     

Effects of liming and boron fertilization on boron uptake of Picea abies

The effects of liming on concentrations of boron and other elements in Norway spruce [Picea abies (L) Karst.] needles and in the mor humus layer were studied in long-term field experiments with and without B fertilizer on podzolic soils in Finland. Liming (2000+4000 kg ha^-1 last applied 12 years before sampling) decreased needle B concentrations in the four youngest needle age classes from 6–10 mg kg^-1 to 5 mg kg^-1. In boron fertilized plots the corresponding concentrations were 23–35 mg kg^-1 in control plots and 21–29 mg kg^-1 in limed plots. Both liming and B fertilizer decreased the Mn concentrations of needles. In the humus layer, total B concentration was increased by both lime and B fertilizer, and Ca and Mg concentrations and pH were still considerably higher in the limed plots than controls. Liming decreased the organic matter concentration in humus layer, whilst B fertilizer increased it.The results about B uptake were confirmed in a pot experiment, in which additionally the roles of increased soil pH and increased soil Ca concentration were separated by means of comparing the effects of CaCO₃ and CaSO₄. Two-year-old bare-rooted Norway spruce seedlings were grown in mor humus during the extension growth of the new shoot. The two doses of lime increased the pH of soil from 4.1 to 5.6 to 6.1, and correspondingly decreased the B concentrations in new needles from 22 to 12 to 9 mg kg^-1. However, CaSO₄ did not affect the pH of the soil or needle B concentrations. Hence the liming effect on boron availability in these soils appeared to be caused by the increased pH rather than increased calcium concentration.     

Relationships between root diameter,root length and root branching along lateral roots in adult,field-grown maize

Background and Aims Root diameter, especially apical diameter, plays an important role in root development and function. The variation in diameter between roots, and along roots, affects root structure and thus the root system’s overall foraging performance. However, the effect of diameter variation on root elongation, branching and topological connections has not been examined systematically in a population of high-order roots, nor along the roots, especially for mature plants grown in the field.Methods A method combining both excavation and analysis was applied to extract and quantify root architectural traits of adult, field-grown maize plants. The relationships between root diameter and other root architectural characteristics are analysed for two maize cultivars.Key Results The basal diameter of the lateral roots (orders 1–3) was highly variable. Basal diameter was partly determined by the diameter of the bearing segment. Basal diameter defined a potential root length, but the lengths of most roots fell far short of this. This was explained partly by differences in the pattern of diameter change along roots. Diameter tended to decrease along most roots, with the steepness of the gradient of decrease depending on basal diameter. The longest roots were those that maintained (or sometimes increased) their diameters during elongation. The branching density (cm^–1) of laterals was also determined by the diameter of the bearing segment. However, the location of this bearing segment along the mother root was also involved – intermediate positions were associated with higher densities of laterals.Conclusions The method used here allows us to obtain very detailed records of the geometry and topology of a complex root system. Basal diameter and the pattern of diameter change along a root were associated with its final length. These relationships are especially useful in simulations of root elongation and branching in source–sink models.     

Differential growth and recovery rates following defoliation in related deciduous and evergreen trees

 Deciduous larches, Larix spp., and evergreen pines, Pinus spp., are sympatric Pinaceae conifers. Adjacent monocultures of 10-year-old Larix decidua Mill. and Pinus resinosa Ait. were subjected to single-season artificial defoliation by clipping from 0% to 99% of each needle. Survival, above-ground productivity, and architecture were measured for 36 months. P. resinosa and L. decidua exhibited differential relationships with defoliation intensity and recovery time. Two months after treatment, defoliation reduced larch height growth but had no effect on radial growth. By contrast, P. resinosa stem radial growth was reduced immediately, but height growth was not decreased until the following year. Pine leader growth and above-ground biomass following 66% defoliation never recovered to control values or 33% defoliated pines. Conversely, defoliated larch quickly recovered from an initial growth loss to eliminate all treatment effects on biomass. The plasticity in architectural response found in larch, but not pine, might partially account for defoliation tolerance. Both P. resinosa and L. decidua exhibited non-linear responses to defoliation. These patterns may be caused partially by the uneven distribution of nutrients within needles, rather than a simple function of leaf area lost to defoliators. Concentrations of 13 nutrients in P. resinosa were highest either in the mid- (Ca, Mg, S, Zn, B, Mn, Fe, Al and Na) or basal- (N, P, K, and Cu) section. The relatively low nutrient content in needle tips may contribute to similar biomass productivity between trees defoliated 33% and controls. Removal of needle mid-sections significantly reduced whole-plant productivity. In contrast, L. decidua nutrients are concentrated in the distal sections. Nutrient concentrations were generally highest in larch. Our results agree with an emergent prediction of the carbon/nutrient balance theory that defoliation more severely reduces growth of evergreen than deciduous species. These results are discussed within the physiological, ecological and evolutionary context of allocation theory, with implications for natural resource management and plant-insect interaction theory. Received: 6 April 1995 / Accepted: 29 August 1995     

Nine Years of Irrigation Cause Vegetation and Fine Root Shifts in a Water-Limited Pine Forest

Scots pines (Pinus sylvestris L.) in the inner-Alpine dry valleys of Switzerland have suffered from increased mortality during the past decades, which has been caused by longer and more frequent dry periods. In addition, a proceeding replacement of Scots pines by pubescent oaks (Quercus pubescens Willd.) has been observed. In 2003, an irrigation experiment was performed to track changes by reducing drought pressure on the natural pine forest. After nine years of irrigation, we observed major adaptations in the vegetation and shifts in Scots pine fine root abundance and structure. Irrigation permitted new plant species to assemble and promote canopy closure with a subsequent loss of herb and moss coverage. Fine root dry weight increased under irrigation and fine roots had a tendency to elongate. Structural composition of fine roots remained unaffected by irrigation, expressing preserved proportions of cellulose, lignin and phenolic substances. A shift to a more negative δ¹³C signal in the fine root C indicates an increased photosynthetic activity in irrigated pine trees. Using radiocarbon (¹⁴C) measurement, a reduced mean age of the fine roots in irrigated plots was revealed. The reason for this is either an increase in newly produced fine roots, supported by the increase in fine root biomass, or a reduced lifespan of fine roots which corresponds to an enhanced turnover rate. Overall, the responses belowground to irrigation are less conspicuous than the more rapid adaptations aboveground. Lagged and conservative adaptations of tree roots with decadal lifespans are challenging to detect, hence demanding for long-term surveys. Investigations concerning fine root turnover rate and degradation processes under a changing climate are crucial for a complete understanding of C cycling.     

THE ROLE OF RESERVES IN LEAVES,BRANCHES, STEMS,AND ROOTS ON SHOOT GROWTH OF RED PINE

By defoliation, girdling, and various combinations of these before the growing season began, the role of foods in old needles, branches, main stems, and roots on shoot growth of 8-yr-old red pine (Pinus resinosa) trees was studied in northern Wisconsin. Defoliation and girdling, alone or in combination, reduced shoot growth but defoliation reduced shoot dry weight more than it reduced shoot elongation. Reductions in shoot growth due to treatment were in the following decreasing order: branch girdling + needle removal > girdling at stem base + needle removal > needle removal > branch girdling > base girdling. The old needles were the major source of food for shoot growth and accounted for four-fifths or more of all shoot growth. The combined reserves in the branches, main stem, and roots accounted for less than 15% of shoot growth. The contribution of reserves from tissues other than old leaves was in the following order: branches > main stem > roots. Defoliation weakened apical dominance relations. In defoliated trees, many secondary axes elongated more than the terminal leader, and secondary axes in lower whorls often grew more than those in upper ones. The data suggest an important role of nutritional factors in correlative growth inhibition.     

NaCl and Na2SO4 alter responses of jack pine (Pinus banksiana) seedlings to boron

Using sand culture, we examined the responses of 6-month-old jack pine (Pinus banksiana Lamb.) seedlings to boron and salinity (sodium chloride and sodium sulfate) treatments. During 4 weeks of treatments, 60 mM NaCl and 60 mM Na₂SO₄ significantly decreased survival, new shoot length, number of new roots, shoot to root dry weight ratio and transpiration rates. When applied in absence of the salts, B had little effect on the measured variables. However, when applied together with salts, B decreased seedling survival, increased needle injury and altered tissue elemental concentrations in jack pine seedlings. In 2 mM B treatment, B concentration was higher in the shoots than in the roots. However, when 2 mM B was present in NaCl and Na₂SO₄ treatments, shoot boron concentration declined and greater proportion of B accumulated in the roots. This shift corresponded to a decline in transpiration rates. In plants treated with NaCl, Na accumulated primarily in the shoots, while in Na₂SO₄-treated plants Na accumulated mostly in the roots. Based on the electrolyte leakage and needle necrosis data, Cl^– appears to be the major factor contributing to seedling injury and B aggravates the injurious effects of NaCl. We suggest that Cl^– may contribute to Na and B toxicity in jack pine by altering cell membrane permeability leading to increased Na concentration in the shoots.     

Root development and configuration in intensively managed radiata pine plantations

Summary In south-east Australia, where radiata pine (Pinus radiata D. Don) is grown on sandy soils low in nutrients and short of water, early establishment, and rapid growth to canopy closure lead to increased productivity. At this stage demands for nutrients and water are high, and trees respond vigorously to silvicultural inputs.For several months after transplanting in winter roots are confined within a narrow planting wedge, low temperature restricts new root growth and slows recovery from water stress in plants. From spring, depending upon the configuration and vigour of the roots transplanted, lateral roots extend radially throughout the soil.Although there were small decreases in concentration of roots radially from the stems of very young trees, such spatial differences disappeared between ages 2 and 3, so that rooting density was independent of distance from the stem. The pattern of vertical distribution of lateral roots was not influenced by age and 80–90% of the lateral roots were within the top 30 cm soil. Roots developed rapidly as the trees grew towards canopy closure, but in general the rooting densities of these pines are among the lowest reported for plants. In rapidly growing trees approaching canopy closure, the secondary thickening of the lateral roots was sufficient to double the weight of roots without altering root length.Knowledge about root growth and root configuration during the early phase of plantation development will assist management decisions where intensive silviculture is practiced, and hence ensure the most efficient use of nutrients and water.     

Traits of Masson Pine Affecting Attack of Pine Wood Nematode

Masson pine characteristics were analyzed in five sample plots in Zhejiang Province, China. Bursaphelenchus xylophilus （Steiner et Buhrer） Nickle （pine wood nematode, PWN） carried by Monochamus alternatus predominately attacked Masson pines in the lower diameter classes. Among the 10 tree characteristics examined, mean crown width, percentage of bole with crown, 5-year cumulative diameter growth, and resin amount showed significant variation between successfully attacked and unattacked trees. The attacked trees had a lower percentage of the bole covered with tree crown, lower crown width, lower radial growth in the last 5 years, and produced less induced resinosis than unattacked trees. Results allowed for effective ranking of the pine forest based on individual tree resistance to PWN. This index of resistance should be considered throughout the development of an ＂Evaluation Criterion and Indicator System＂. The preceding ranking can be used to evaluate the resistance and resiliency of the pine forest ecosystem to PWN＇s invasion, which is similar to Pest Risk Analysis （PRA）.