Determinants of woody species richness in Scot pine and beech forests: climate, forest patch size and forest structure

We analysed patterns of woody species richness in Pinus sylvestris and Fagus sylvatica forests in Catalonia (NE Spain) from forestry inventory databank in relation to climate and landscape structure. Both types of forests are found within the same climatic range, although they have been managed following somewhat different goals. Overall, woody species richness significantly increased when conditions get closer to the Mediterranean ones, with milder temperatures. Differences between the two types of forests arose when comparing the relationship between richness and forest patch size. Woody species richness increased in pine forests with patch size, while the opposite trend was observed in beech forests. This pattern is explained by the different behaviour of structural canopy properties, since leaf area index and canopy cover showed a steeper increase with increasing forest patch size in Fagus forests than in Pinus ones. Accordingly, richness decreased with canopy cover in Fagus plots, but not in Pinus ones. We suggest that these differences would be related to management history, which may have enhanced the preservation of beech stands in larger forest landscape units.     

Chemical composition and ultrastructural changes in Scots pine needles in a forest decline area in southwestern Finland

Tree decline and deaths have been observed among 15 to 20-year-old Scots pines (Pinus sylvestris L.) in a dry heath forest in southwestern Finland. The sudden decline in height growth, the dieback of leading shoots and the yellowing of needles in young shoots in the upper part of the tree are typical symptoms of the decline of these young pines. Needle ultrastructure and chemical composition of Scots pines with or without decline and fluctuations of them in different seasons were studied. Afflicted trees were found to suffer from a deficiency in calcium and magnesium with low concentration of foliar nitrogen and phosphorus observed in all the trees studied. Ultrastructural study revealed changes characteristic of different seasons and measured nutrient status of needles. A clear reduction of membrane system in chloroplasts, especially related to Mg deficiency, was observed in most samples. The symptoms related to N deficiency, the translucent appearance of the cytoplasm and chloroplast stroma, and the elongated chloroplasts, as well as swelling of mitochondria, indicating P deficiency, were also found in the needles sampled from this forest decline area. The present study showed that it is possible to detect specific nutrient deficiency symptoms in needle ultrastructure in field samples and for use as sensitive indicators of unbalanced nutrient status.     

沙地樟子松天然林南缘分布区林木竞争、空间格局及其更新特征

以呼伦贝尔沙地樟子松地理分布南缘天然林为研究对象,采用Hegyi竞争指数与点格局等分析方法,研究了沙地樟子松天然林的种群结构、种内竞争、空间格局及幼树更新等特征。结果表明:1)所调查区域天然沙地樟子松纯林处于中幼龄阶段,属增长型种群;2)其竞争指数与对象木胸径服从幂函数关系CI=242.24D~(-1.12)(R~2=0.91);3)幼树在小尺度上呈聚集分布,中树与大树在中大尺度上呈随机分布;幼树与中树在小尺度上呈正相关性,中树与大树在小尺度呈负相关性;4)竞争指数与更新幼树和存活更新幼树的密度均呈显著正相关性。在林分管理中需要充分考虑林木竞争、空间格局以及种群更新的关系,本研究可为沙地樟子松天然林的经营管理与保护提供重要的科学依据。     

Resistance of Scots pine wood to Brown-rot fungi after long-term forest fertilization

The susceptibility of Scots pine (Pinus sylvestris L.) sap- and heartwood against the wood decaying brown-rot fungus (Coniophora puteana) was investigated after long-term forest fertilization at three different sites in central Finland. Different wood properties: wood extractives, wood chemistry, and wood anatomy were used to explain sap- and heartwood decay. Scots pine sapwood was more susceptible to decay than its heartwood. In one site, sapwood seemed to be more resistant to wood decay after forest fertilization whereas the susceptibility of heartwood increased. Significant changes in the sapwood chemistry were found between treatment and sites, however, no relationship between wood chemistry and wood decay was observed in the factor analysis. The results of this study show that there was an inconsistent relationship between decay susceptibility and fertilization and the measured physical and chemical attributes of the wood were not consistently correlated with the decay rate.     

樟子松固沙林生长对土壤磷素变化的影响

磷是森林生态系统重要养分元素之一,是干旱半干旱地区植物生长的限制性因子。然而沙质草地转化为人工林生态系统后,林分的生长对沙地土壤磷素的变化及其影响机理还不清晰,为沙地合理经营和管理人工林带来了不确定性。以辽宁省章古台地区各生长阶段(幼龄林、中龄林、成熟林和过熟林阶段)的20块樟子松固沙林样地(在各生长阶段林分附近寻找1块天然草地作为对照样地)为研究对象,取样并测定样地各土层(0—10、10—20、20—40、40—60、60—80、80—100 cm)的土壤磷(全磷和速效磷)、土壤氮(全氮和速效氮)、土壤钾(全钾及速效钾)、土壤有机碳等含量以及土壤含水率、土壤pH值、土壤质地、土壤容重等因子值,并进行统计分析。结果表明:沙质草地营造樟子松人工林后,土壤全磷含量随林龄的增加而逐渐递增,成熟林时期达到最高,过熟林地全磷降低,且土壤全磷对土层深度不敏感。成熟林地的速效磷含量略高于幼林和中龄林;虽然与幼林、中龄林没有显著差异,但过熟林的土壤速效磷含量是所有林分中最低的。除了林分生长影响外,草地营造樟子松林后,土壤全磷的变化还受土壤容重和土壤速效氮含量的影响,而土壤速效磷的变化受土壤有机碳和pH...     

Spermidine and the ectomycorrhizal fungus Pisolithus tinctorius synergistically induce maturation of Scots pine embryogenic cultures

Exogenous spermidine (Spd) and the ectomycorrhizal (ECM) fungus Pisolithus tinctorius (Pers.) Coker and Couch had a synergistic effect on the maturation of Scots pine (Pinus sylvestris L.) somatic embryos. Induced maturation was expressed as a higher number of cell masses able to form embryos and a greater number of embryos formed per cell mass. In contrast, treatment with P. tinctorius alone on the hormone-free medium resulted in the lowest embryo-forming capacity. Retarded proliferation growth appeared to be required for maturation, but did not explain the synergistic effect of the fungus and exogenous Spd. Simultaneous treatment did not result in lower concentrations of putrescine (Put), Spd or spermine (Spm) in the embryogenic cell masses relative to the separate treatments. Our study is the first report on the use of a specific ECM fungus to induce maturation of somatic embryos, and it indicates that P. tinctorius was able to modify the maturation media in a way that, together with exogenous Spd, positively affected embryogenic cultures of Scots pine. Our study also shows that it is possible to enhance plant development other than root formation by using specific ECM fungi.     

Post-fire regeneration in a Mediterranean pine forest with historically low fire frequency

Species of Mediterranean vegetation are known to regenerate directly after fire. The phenomenon of autosuccession (direct regeneration) has been found to be often combined with an increase of species richness during the first years after fire due to the high abundance of short-lived herbaceous plants facilitated by plentiful nutrients and light. The high degree of vegetation resilience, which is expressed in terms of autosuccession, has been explained by the selective pressure of fire in historic times. According to existing palaeoecological data, however, the Pinus halepensis forests in the Ricote Mountains (Province of Murcia, SE Spain) did not experience substantial fire impact before the presence of man nor are they especially fire-prone today. Therefore, we studied post-fire regeneration to find out if direct succession is present or if species from pre-fire vegetation are absent during the post-fire regeneration stages. Patterns of succession were deduced from observations made in sample plots on sites of a known regeneration age as well as in adjacent unburnt areas. The results of the vegetation analyses, including a Detrended Correspondence Analysis, indicate that Pinus halepensis forest regeneration after fire resembles autosuccession. As regards the presence of woody species, there is a high percentage similarity on north (83%) and south (70%) facing slopes during the first year after fire vs. reference areas which is due, for example, to direct regeneration of the resprouting Quercus coccifera or seeders like Pinus halepensis or Fumana laevipes. However, if herbaceous species are included in the comparison, the similarity on north-facing sites decreases (to 53%) with the presence of additional species, mainly ruderals like Anagallis arvensis or Reseda phyteuma, and even woody species on the burnt plots. This effect indicates “enhanced autosuccession”, which was not found on south-facing sites where overall species richness was very high irrespective of the impact of fire. Locally we found limited regeneration of some species, for example Pinus halepensis at high altitudes (1000 m), even 22 years after fire. As we assume that historical fires did not play an important role in the area and direct succession is present nevertheless, our results support the theory that autosuccession is not a process restricted to fire-prone areas. Fire has been only one of several selective forces since human settlement that probably led to a set of species pre-adapted against recurrent disturbance.     

Seed colour and post-fire bird predation in a Mediterranean pine forest

In a Pinus halepensis Mill. forest, a field experiment was designed to evaluate post-fire seed predation as affected by combinations of seed colour and soil substrates: light grey and black seeds combined with light grey ash, dark grey ash and pale brown sand. A survey of bird species inhabiting the area was also carried out and polyphenolic content of seed coat was assessed in seed lots of different colour. Light grey seeds were observed to be less predated on light grey ash, suggesting eucrypsis as a protective strategy against bird predation. On the contrary, no clear pattern was observed for the predation of black seeds on different substrates. In the study area both bird species breaking the seed coat and eating the endosperm and bird species swallowing the whole seed were monitored. We have estimated that more seeds were swallowed than broken, in all colour categories. Light grey seeds, which were found to have a higher content of polyphenols, were predated more than black seeds when exposed on the same substrate. Thus, no evidence was produced that the amount of polyphenols in seed coat could protect seeds from predation.     

Nutritional disturbances of young Scots pines caused by pine bark bugs in a dry heath forest

This study deals with the effect of pine bark bugs (Aradus cinnamomeus Panzer) on the nutrition of young Scots pines (Pinus sylvestris L.). Soil and needle samples for analytical purposes were collected from a young pine stand growing on a dry barren mineral soil afflicted by pine bark bugs.The damage to vascular tissues caused by pine bark bugs disturbed the nutrition of the trees, especially in the top part of the crown. The foliar calcium, magnesium, manganese and sulphur concentrations were highly reduced. Scots pines suffered from a lack of calcium, magnesium, nitrogen and phosphorus. These deficiencies were secondary by nature, because no differences were observed between the nutrient concentrations of the underlying soil of the healthy and affected trees. The symptoms of trees damaged by pine bark bugs resembled most of all calcium deficiency symptoms.     

Microscopic structure of Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> (L.)) needles during ageing and autumnal senescence

Needle ageing and senescence were studied in Scots pine (Pinus sylvestris) trees growing in natural conditions with minimal anthropogenic influence. The four existing needle generations were analyzed by light and transmission electron microscopy before and during autumnal yellowing of the oldest needle generation. The change from green to yellow occurred within less than 4 days in central Finland. The structure of the oldest needles remained largely intact as long as they were green. Increase in mitochondrion size and hypertrophy of the phloem parenchyma were the only changes, probably related to the approaching senescence. In the yellow needles, the structure of the mesophyll tissue varied from nearly intact with reduced chloroplasts and higher numbers of plastoglobuli, to totally disintegrated cells. In the disintegrated cells, peroxisomes were absent, and chloroplasts were smaller with a patchy appearance and degraded, eventually empty-looking stroma. Mitochondria were enlarged, but retained integrity until the last stages of deterioration, and lipids increased. At the light microscopic level, vacuolar volume in mesophyll cells and cavity formation in transfusion tissue increased. Ageing was characterized by increases in the vacuolar volume and cytoplasmic lipids, altered appearance of vacuolar tannin from homogenous ‘sandy’, to large spherical drops and finally to a large mass in the mesophyll, and by hypertrophy and tannin accumulation in the phloem parenchyma. Changes related to needle ageing, senescence and cell location in the mesophyll tissue were discussed relative to findings with stress by strong light, weather conditions and ozone.     

Spermidine and methylglyoxal bis(guanylhydrazone) affect maturation and endogenous polyamine content of Scots pine embryogenic cultures

Exogenous spermidine (Spd) and methylglyoxal bis(guanylhydrazone) (MGBG), a putative inhibitor of Spd synthesis, improved somatic embryo formation of Scots pine (Pinus sylvestris L.). The induced maturation due to MGBG and Spd was accompanied by significantly retarded proliferation growth and by reduction in the concentration of free polyamines compared to the control cultures. The action of MGBG revealed that it has a non-specific effect on the whole polyamine metabolism of Scots pine. Furthermore, at certain concentrations it may induce plant differentiation as well.     

Lignosulfonate promotes the interaction between Scots pine and an ectomycorrhizal fungus Pisolithus tinctorius in vitro

Lignosulfonate (LS) is a lignin-based polymer obtained as a by-product from paper industry, which may have potential as an amendment with macronutrients. We studied effects of LS on the interaction between Scots pine (Pinus sylvestris L.) seedlings and hypocotyl cuttings and the ectomycorrhizal (ECM) fungusPisolithus tinctorius (Pers.) Coker and Couch. The experiments were performed in vitroon the MMN agar medium containing Fe–LS chelate at the concentrations of 0, 5, 10 and 25 mg/L. Inoculation with P. tinctoriusincreased root growth of the seedlings. Fe–LS enhanced P. tinctorius induced formation of lateral roots and had a dose-dependent positive effect on the establishment of mycorrhizas on the seedlings. The growth of the fungal mycelium was improved by Fe–LS, which might cause faster and more intensive contact with the roots and, thus, better root growth and mycorrhiza formation. P.tinctorius enhanced also adventitious root formation and subsequent root growth of the hypocotyl cuttings but without any synergistic effect with Fe–LS. Our study with P. tinctorius and Scots pine in vitro indicates that a low-cost by-product Fe–LS, obtained from paper industry, may be a potential tool to improve the efficiency of fungal inoculations, thus, facilitating the early interaction between an ECM fungus and host seedling.     

Recovery from winter depression of photosynthesis in pine and spruce

Summary Recovery from winter depression of photosynthesis was studied in Pinus sylvestris, Pinus conforta and Picea abies by means of chlorophyll fluorescence and gas exchange measurements. During the winter 1986–1987 the fluorescence yield was low and no variable fluorescence was detectable before the end of March. In the field recovery of variable fluorescence/maximum fluorescence (F_v/F_m) during spring was slow for all three species studied. The temperature dependence of recovery was confirmed from measurements of the potential rate of recovery of F_v/F_m at different temperatures in the laboratory. At 20° C, F_v/F_m increased from 0.1 to 0.8 within 3 days. Recovery of F_v/F_m was paralleled by an increase in apparent photon yield. No significant differences could be demonstrated between the studied tree species in potential rate of recovery in the laboratory or in actual recovery in the field.     

Long-term needle litterfall of a Scots pine Pinus sylvestris stand: relation to temperature factors

Summary Needle litterfall of a Scots pine was caught over 24 years (1962–1986) with litter-traps in a Scots pine stand in southeastern Finland. The age of the trees averaged 111 years in 1962. The stand was naturally recruited and only minor silvicultural treatments occurred during its history. Litterfall showed great year-to-year variation, the minimum being 18 g/m² (in 1968) and maximum 213 g/m² (in 1973). There was no overall trend in the amount of litterfall, and the age of the stand was thus not important in determining the needle fall. We used time domain time series analysis (ARIMA) and standard climatic data (temperature, precipitation) to investigate the relationship of litterfall to climatic factors. Mean July temperature was clearly correlated with needle litterfall. High temperature in July coincided with enhanced litterfall in the same and the next year. Litterfall enhanced litterfall in the same and the next year. Litterfall increased also after high temperatures during March–April, but only in the same year. In addition to these the litterfall had a 4-year self-dependency. This is approximately the same as the mean longevity of needles in the study area. Altogether the time series model we propose covers about 90% of the variance of the original time series.     

The ancient use ofPinus sylvestris L. (Scots pine) inner bark by Sami people in northern Sweden, related to cultural and ecological factors

A multidisciplinary study of use by Sami people ofPinus sylvestris L. (Scots pine) inner bark was performed in northern Sweden. We combined linguistic, historical and archaeological records with ecological data collected from field studies to investigate important cultural and ecological factors associated with previous use of bark. Our data from bark peeled trees at 313 sites were used to describe patterns in space and time in the land-scape. Sites with bark peeled trees were mainly found in the central and northern part of Lapland (65°00–68°14N). Large numbers of barked trees were found in undisturbed forests in national parks and reserves but few were found in forests under commercial management. The dendroecological analysis revealed a continuous use of inner bark from A.D. 1450 to 1890. Large sheets of bark were taken from trees in the spring, prepared and stored as a staple food resource. Inner bark was eaten fresh, dried or roasted. Smaller bark peelings were used for the wrapping of sinews. The cessation of bark use in the 19th century was driven by several factors, but the availability of other products that could replace traditional use of bark was specifically important. The previous common and varied use of bark, the great age of Sami terminology and a possible association with archaeological remains of potential great age indicate that peeled bark was used long before the historic period. Our study also focused on the ongoing loss of culturally-modified trees in forests outside protected areas.     

Freeze-preservation of buds from Scots pine trees

A procedure has been developed for freeze-preservation of buds of the Scots pine (Pinus sylvestris L.). Instead of liquid nitrogen, cold storage in –80°C was used. The partly dormant material used in the experiments was obtained directly from a natural stand in Northern Finland and no prefreezing or cryoprotectants for preconditioning were used. Cooling velocity was 1°C/min up to a terminal freezing temperature of –39°C, after which the buds were immersed in liquid nitrogen at –196°C for 10 minutes. The material was then transferred to a deepfreezer at –80°C and stored up to 6 months. After rapid thawing, the buds were sterilized and their viability was tested by FDA staining and by culturing meristems on 1/2 MS medium for at least two weeks. All the freezing experiments were performed during March and April. The best survival of buds (90–100%) was achieved at the beginning of April, after which a pronounced decline in survival occurred obviously due to a rise in the water content of the buds.     

Leaf and canopy responses to elevated CO2 in a pine forest under free-air CO2 enrichment

Physiological responses to elevated CO₂ at the leaf and canopy-level were studied in an intact pine (Pinus taeda) forest ecosystem exposed to elevated CO₂ using a free-air CO₂ enrichment (FACE) technique. Normalized canopy water-use of trees exposed to elevated CO₂ over an 8-day exposure period was similar to that of trees exposed to current ambient CO₂ under sunny conditions. During a portion of the exposure period when sky conditions were cloudy, CO₂-exposed trees showed minor (7%) but significant reductions in relative sap flux density compared to trees under ambient CO₂ conditions. Short-term (minutes) direct stomatal responses to elevated CO₂ were also relatively weak (5% reduction in stomatal aperture in response to high CO₂ concentrations). We observed no evidence of adjustment in stomatal conductance in foliage grown under elevated CO₂ for nearly 80 days compared to foliage grown under current ambient CO₂, so intrinsic leaf water-use efficiency at elevated CO₂ was enhanced primarily by direct responses of photosynthesis to CO₂. We did not detect statistical differences in parameters from photosynthetic responses to intercellular CO₂ (A _net-C _i curves) for Pinus taeda foliage grown under elevated CO₂ (550 mol mol^–1) for 50–80 days compared to those for foliage grown under current ambient CO₂ from similar-sized reference trees nearby. In both cases, leaf net photosynthetic rate at 550 mol mol^–1 CO₂ was enhanced by approximately 65% compared to the rate at ambient CO₂ (350 mol mol^–1). A similar level of enhancement under elevated CO₂ was observed for daily photosynthesis under field conditions on a sunny day. While enhancement of photosynthesis by elevated CO₂ during the study period appears to be primarily attributable to direct photosynthetic responses to CO₂ in the pine forest, longer-term CO₂ responses and feedbacks remain to be evaluated.     

Fine root branch orders respond differentially to carbon source-sink manipulations in a longleaf pine forest

Fine roots are a key component of carbon (C) flow and nitrogen (N) cycling in forest ecosystems. However, the complexity and heterogeneity of the fine root branching system have hampered the assessment and prediction of C and N dynamics at ecosystem scales. We examined how root morphology, biomass, and chemistry differed with root branch orders (1–5 with root tips classified as first order roots) and how different root orders responded to increased C sink strength (via N fertilization) and reduced carbon source strength (via canopy scorching) in a longleaf pine (Pinus palustris L.) ecosystem. With increasing root order, the diameter and length of individual roots increased, whereas the specific root length decreased. Total root biomass on an areal basis was similar among the first four orders but increased for the fifth order roots. Consequently, total root length and total root surface area decreased systematically with increasing root order. Fine root N and lignin concentrations decreased, while total non-structural carbohydrate (TNC) and cellulose concentrations increased with increasing root order. N addition and canopy disturbance did not alter root morphology, but they did influence root chemistry. N fertilization increased fine root N concentration and content per unit area in all five orders, while canopy scorching decreased root N concentration. Moreover, TNC concentration and content in fifth order roots were also reduced by canopy scorching. Our results indicate that the small, fragile, and more easily overlooked first and second order roots may be disproportionately important in ecosystem scale C and N fluxes due to their large proportions of fine root biomass, high N concentrations, relatively short lifespans, and potentially high decomposition rates.Electronic Supplementary Material Supplementary material is available for this article at     

Nitrogen mineralization and H+ transfers in a Scots pine (Pinus sylvestris L.) forest soil as affected by liming

H⁺ production due to N uptake in a mature Scots pine stand subjected to high NH₄ ⁺ deposition was previously estimated to amount to approx. 2.2 kmol ha^-1 y^-1. The question whether H⁺ transfers related to N mineralization (ammonification and nitrification) offset or corroborate this proton production is investigated in the present research. To determine N mineralization, soil cores were used of which both ends were closed with layers of ion exchange resin (IER) to prevent influx and efflux of ions. The effect of liming on N mineralization and the resulting H⁺ production was investigated in 7 incubation periods of each ca. 8 wk. Because of its high mobility NO₃ accumulated in both IER layers at the expense of that in the incubated forest floor and mineral soil. Net N mineralization in the soil cores as a whole amounted to 40 and 77 kg N ha^-1 in 384 d in the control and limed plots, respectively. In both treatments ca. 65% of mineralized N was nitrified. H⁺ production due to N mineralization amounted to approx. 1.2 kmol ha^-1 y^-1 in the control and limed plots. Liming reduced the amount of C in the forest floor, but not forest floor mass, because of an increased mixing with mineral particles.