Harvest impacts on soil carbon storage in temperate forests

Forest soil carbon (C) storage is a significant component of the global C cycle, and is important for sustaining forest productivity. Although forest management may have substantial impacts on soil C storage, experimental data from forest harvesting studies have not been synthesized recently. To quantify the effects of harvesting on soil C, and to identify sources of variation in soil C responses to harvest, we used meta-analysis to test a database of 432 soil C response ratios drawn from temperate forest harvest studies around the world. Harvesting reduced soil C by an average of 8 ± 3% (95% CI), although numerous sources of variation mediated this significant, overall effect. In particular, we found that C concentrations and C pool sizes responded differently to harvesting, and forest floors were more likely to lose C than mineral soils. Harvesting caused forest floor C storage to decline by a remarkably consistent 30 ± 6%, but losses were significantly smaller in coniferous/mixed stands (−20%) than hardwoods (−36%). Mineral soils showed no significant, overall change in C storage due to harvest, and variation among mineral soils was best explained by soil taxonomy. Alfisols and Spodosols exhibited no significant changes, and Inceptisols and Ultisols lost mineral soil C (−13% and −7%, respectively). However, these C losses were neither permanent nor unavoidable. Controls on variation within orders were not consistent, but included species composition, time, and sampling depth. Temporal patterns and soil C budgets suggest that forest floor C losses probably have a lesser impact on total soil C storage on Alfisols, Inceptisols, and Ultisols than on Spodosols, which store proportionately large amounts of C in forest floors with long C recovery times (50–70 years). Mineral soil C losses on Inceptisols and Ultisols indicate that these orders are vulnerable to significant harvest-induced changes in total soil C storage, but alternative residue management and site preparation techniques, and the passage of time, may mitigate or negate these losses. Key findings of this analysis, including the dependence of forest floor and mineral soil C storage changes on species composition and soil taxonomic order, suggest that further primary research may make it possible to create predictive maps of forest harvesting effects on soil C storage.     

Liana habitat and host preferences in northern temperate forests

Lianas and other climbers are important ecological and structural components of forest communities. Like other plants, their abundance in a given habitat depends on a variety of factors, such as light, soil moisture and nutrients. However, since lianas require external support, host tree characteristics also influence their distribution. Lianas are conspicuous life forms in tropical regions, but in temperate areas, where they are less prominent, little is known about factors that control their distributions in these forests. We surveyed the climbing plant species in 20 mature (100 years and greater) forested habitats in the Midwest USA at a variety of levels from simple presence/absence, to ground layer abundances, to those species that had ascended trees. We also examined attributes of the tree species with climbers attached to them. Using cluster analysis, we distinguished five different tree communities in our survey locations. We determined that 25% of the trees we surveyed had one or more lianas attached to it, with Parthenocissus quinquefolia (Virginia creeper) the most common climbing species encountered. Canopy cover and soil attributes both influenced climber species presence/absence and ground layer climber abundance. The proportion of liana species of a given climbing type (roots, stem twiner, tendril climber) was significantly related to the DBH of the host tree, with more root climbers and fewer stem and tendril climbers on large trees. In general, the DBH of climbing lianas had a significant positive relationship to the DBH of the host tree; however this varied by the identity of the liana and the tree species. The greater the DBH of the host tree, the higher the probability that it was colonized by one or more lianas, with tree species such as Pinus banksiana (jack pine) and Quercus alba (white oak) being more susceptible to liana colonization than others. Finally, some liana species such as Celastrus scandens (American bittersweet) showed a preference for certain tree species (i.e., P. banksiana) as hosts. The information obtained about the relationship between the tree and climber community in this study provides insight into some of the factors that influence liana distributions in understudied temperate forest habitats and how lianas contribute to the structure of these mature forests. In addition, these data can provide a point of comparison to other liana communities in both temperate and tropical regions.     

Controls of growth phenology vary in seedlings of three, co-occurring ecologically distinct northern conifers

The objective of this study was to investigate the effects of temperature and seed-source elevation on height-growth phenology of three co-occurring and ecologically distinct northern conifers (Pinus contorta Dougl. ex Loud. var. latifolia (lodgepole pine), Picea glauca (Moench) Voss x Picea engelmannii Parry ex Engelm. (interior spruce) and Abies lasiocarpa (Hook.) Nutt. (subalpine fir)). Seed from populations of the three indigenous and co-occurring species was collected across an elevational transect on the southwestern slope of McBride Peak, near Prince George, BC. Collection sites were at elevations of 750 to 1850 m, the latter being close to the tree line. In 2003, seeds were germinated and seedlings raised under favorable growing conditions in a temperature-controlled glasshouse. In 2004, seedlings of each population were grown in natural daylengths at a location within 50 km of the seed collection site both in a temperature-controlled glasshouse and at a nearby field site, and height growth was recorded twice a week throughout the growing season. Species differed in both the date and the accumulated heat sum above 5 degrees C for the initiation and cessation of shoot extension. Growth durations (which integrate growth initiation and growth cessation) were more similar among species in the field than in the glasshouse. This suggests that different mechanisms of phenological control among co-occurring species can result in adaptive "equivalence" under a particular set of climatic conditions.     

冀北地区华北落叶松人工林提前主伐利用的探讨

通过对冀北地区围场县落叶松人工林的调查,从落叶松的林龄结构、落叶松采伐的经济效益、落叶松适生的立地条件及提前主伐利用对非公有制林业的激励等方面,论述了冀北地区华北落叶松提前主伐利用在促进林业发展、发挥生态作用的可行性和重要意义。     

Relationships among drought resistance, transpiration rates, and abscisic acid levels in three northern conifers

Water potentials, transpiration rates and abscisic acid (ABA) levels in shoots of black spruce (Picea mariana (Mill.) BSP), white spruce (Picea glauca (Moench) Voss) and jack pine (Pinus banksiana Lamb.) seedlings were monitored during periods of drought and recovery from drought. Abscisic acid contents of shoots increased during the period of drought as water potentials decreased. The increase in levels of ABA was closely associated with a decrease in rates of transpiration. In the spruces, the levels of ABA peaked and then fell while plant water potentials continued to decrease, whereas in jack pine, the level of ABA rose throughout the drought treatment. After rewatering, the levels of ABA in all three conifers fell concurrent with a rise in transpiration rates. At the end of the three-day recovery period, ABA levels and transpiration rates in the spruces were either at or near control levels, whereas the concentration of ABA in jack pine remained approximately twice the control level, and transpiration was only 60% of the control rate. A compound tentatively identified as phaseic acid followed trends similar to those for ABA.     

Accumulation of free amino acids in the shoots and roots of three northern conifers during drought

The effects of drought on the free amino acid pools in 21- to 23-week-old seedlings of black spruce (Picea mariana (Mill.) Britt.), white spruce (Picea glauca (Moench.) Voss) and jack pine (Pinus banksiana Lamb.) were followed during soil drying. Although water and pressure potentials were sensitive to water deficits, large changes in osmotic potential were not recorded until after the development of severe drought. Total soluble amino nitrogen in the shoots and roots of the three species rose as turgor declined, with peak concentrations attained late in the drought period when the pressure potentials of the shoots approached zero. All white spruce seedlings were alive at zero turgor and showed large decrements in osmotic potential, but concentrations of free amino nitrogen in the roots and shoots showed only modest increases, reaching 125 to 150% of their control values. In contrast, large numbers of black spruce and jack pine were dead or severely damaged at zero turgor, and only small changes in osmotic potential were detected during soil drying. Nevertheless, concentrations of soluble amino nitrogen in both species reached 150 to 200% of control values a few days before the seedlings died. Alanine, arginine, aspartic acid/asparagine, glutamic acid/glutamine, glycine, hydroxyproline and proline were the major components of the free amino acid pools under both water-stressed and non-stressed conditions, with the largest and most consistent increases observed in the roots of all three conifers. Although proline was an important and dynamic component of the free pools, absolute concentrations were commony equalled or exceeded by other free amino acids in the roots and shoots and nearly always exceeded by the concentration of aspartic acid/asparagine in both tissues. Differences in drought resistance among the three conifers were not reflected by unique patterns of amino acid accumulation or by large differences in absolute concentrations of the free amino acid pools.     

Long-term charcoal-induced changes to soil properties in temperate regions of northern Iran

The long-term performance and benefits of charcoal application on the carbon sequestration and properties of forest soils in temperate or non-tropical regions has not been studied in detail in spite of its important role in global warming. This study was conducted to describe the long-term charcoal-induced changes in organic carbon (OC) content and other soil properties of temperate deciduous forests in Mazandaran province, northern Iran. Three sites were sampled to collect composite soil samples from two depths (0–20 and 20–40 cm) inside and outside of a plot of charcoal-enriched soils surrounding a historical charcoal production site (abandoned for more than 120 years). The presence of charcoal in soils for about 120 years elevated significantly the black carbon, total OC, natural soil OC, total nitrogen, dissolved organic matter, soil OC density, exchangeable bases, saturated hydraulic conductivity, available water capacity and available Fe, Mn and Zn compared to the adjacent reference soils. Cation exchange capacity (CEC) and pH were 15.5 cmolc kg^-1 and 0.5 units, respectively, higher than the adjacent reference soils at 0–20 cm soil depth. However, electrical conductivity (EC), bulk density and available Cu were higher in the adjacent reference soil. The aged charcoal had no significant effect on the microbial respiration rate of studied soils. The results of this study provide new insights and strong support for the long-term benefits of biochar application as a management strategy for improving soil productivity as well as sequestering large quantities of durable carbon in soils of the region and mitigating global warming.     

First frost: Effects of single and repeated freezing events on acclimation in Picea abies and other boreal and temperate conifers

In experiments with needles of Picea abies, we tested the specific hypothesis that a single night of freezing acts as a signal that triggers a rapid increase in low temperature (LT) tolerance, and the more general hypothesis that repeated or prolonged freezing stimulates increased LT acclimation. In three growth chamber experiments involving acclimation under early- to mid-autumn light and temperature conditions followed by one or more freezing treatments, we found no significant effect of a single night of freezing on LT tolerance, and only limited and inconsistent effects of repeated and prolonged freezing. We also tested the effect of prolonged storage at −5 °C on LT tolerance on samples of three boreal and three temperate conifer species during acclimation under field conditions, and again found no consistent enhancement of LT tolerance attributable to freezing in either group. In agreement with our own and others’ anecdotal observations that some species can attain nearly maximal LT tolerance in the absence of freezing under field conditions, we conclude that freezing is neither required nor a major influence in LT acclimation, at least in well-studied boreal conifer species, while the effects of freezing on temperate conifers are not as well-documented. We conclude that freezing treatment of conifer seedlings to ensure sufficient hardiness for late planting seems to offer little practical advantage.     

Foliage browning and shoot death in Sitka spruce and other conifers in northern Britain during winter 1983–84

A hitherto undescribed form of injury to Sitka spruce and other conifers occurred in plantations throughout northern Britain during the dormant season 1983–84. Damage consisted primarily of shoot death which was typically concentrated in a zone of variable extent below the upper few whorls of the tree. It occurred predominantly in forests in the west and was most severe in older crops at high elevation. Possible causes are discussed and it is concluded that the damage was probably climatic in origin.     

Factors affecting somatic embryogenesis in conifers

This review seeks to examine the extreme response of isolated somatic plant cells of apical meristematic tissues of mature conifer trees towards specific stress conditions in vitro resulting in somatic embryogenesis.Signal molecules regulating embryo development have been described in angiosperms,but very little is known about somatic rejuvenation in conifers.Recent studies on cloning of mature conifers provide new perspectives on signal molecules on cellular dedifferentiation into the embryogenic pathway.Our recent studies show that signal molecules such as butenolide,calcium ions,salicylic acid,antioxidants,amino acids,triacontanol and 24-epibrassinolide all play an important role in the conversion of somatic cells into an embryogenic pathway in many recalcitrant pines.This constitutes a major breakthrough in forest biotechnology with many practical applications in clonal forestry.     

Coarse woody debris features of a warm temperate deciduous broad-leaved forest,northern China

Stocking and structural composition of a deciduous broad-leaved forest were determined to predict coarse woody debris quantity by quantifying the empirical relationships between these two attributes. The most ecologically significant families by stem density were Salicaceae, Betulaceae, Fagaceae, and Aceraceae. P opulus davidiana was the most dominant species followed by B etula dahurica, Quercus mongolica, and Acer mono. The four species accounted for 69.5% of total stems. Numerous small-diameter species characterized the coarse woody debris showing a reversed J-shaped distribution. The coarse debris of P. davidiana, B. dahurica, and Q. mongolica mainly comprised the 10–20 cm size class, whereas A. mono debris was mainly in the 5–10 cm size class. The spatial patterns of different size classes of coarse woody debris were analyzed using the g-function to determine the size of the tree at its death. The results indicate that the spatial patterns at the 0–50 m scale shifted gradually from an aggregated to a random pattern. For some species, the larger coarse debris might change from an aggregated to a random distribution more easily. Given the importance of coarse woody debris in forest ecosystems, its composition and patterns can improve understanding of community structure and dynamics. The aggregation pattern might be due to density dependence and self-thinning effects, as well as by succession and mortality. The four dominant species across the different size classes showed distinct aggregated distribution features at different spatial scales. This suggests a correlation between the dominant species population, size class, and aggregated distribution of coarse woody debris.     

Stump infection by Armillaria in first-rotation conifers

Information about the entry of Armillaria into first-rotation pine and spruce stands was obtained by searching for infected stumps, rhizomorph systems or trees that had been killed. In pines Armillaria foci were very rare. In pure Norway spruce Armillaria lutea and A. mellea were detected in stumps but rhizomorphs did not extend into the soil; in Norway spruce mixed with oak, by contrast, A. lutea sometimes produced extensive rhizomorph systems. In Sitka spruce small groups of trees had been killed by A. ostoyae. All foci investigated in conifers contained different genotypes of Armillaria and probably originated from spore infection of stumps created by thinning. Some implications of these findings are discussed.     

The silviculture of conifers in Great Britain

The original paper by H.M. Steven, with this title, describedthe silvicultural characteristics of the main coniferous speciesthen available in British forestry. The use of these introducedconifers was justified by the limited site tolerances of theone native species, Scots pine, and the need to expand forestcover on to sites with infertile soils in harsh climatic conditions. This paper reviews the developments in silvicultural techniquesand understanding that have enabled the rapid expansion of productiveforestry in the last seventy years:

• The problems associated with afforestation, requiring ameliorationof soil physical and chemical conditions, have been resolvedand the climatic limits to the use of individual species aremore clearly defined.
• Increased knowledge of the physiologicalresponses of individualspecies to environmental factors hasinfluenced silviculturalpractice from nursery production tothe regeneration of maturestands.
• Analysis of genetic variationwithin introduced conifer specieshas refined their use andis leading to improvements in theirproductivity and timberquality.
• The application of ecological principles, based onthe ecosystemconcept, has led to an understanding of foreststand dynamicsthat should ensure production from exotic specieson a sustainablebasis.

Future requirements in the silviculture of conifers includethe transferof the results of computer simulation modellinginto practice, the developmentof silvicultural systems appropriateto British conditions and greater emphasis on the quality ofproduction.     

Nematological problems in the production of conifers

Several conifer nurseries were visited in Piedmont (Italy) and soil and root-samples were taken and examined for plant-parasitic nematodes. In these investigations damage to the mycorrhizae caused by phytohelminths, most frequently through their trophic activity on the tissue of the host plant, has been observed.     

Generalized functions of biomass expansion factors for conifers and broadleaved by stand age,growing stock and site index

Parties to the Kyoto Protocol and/or the United Nations Framework Convention on Climate Change (UNFCCC) are required to account for their direct human-induced carbon emissions and removals including those from forestry and other land use related activities. In most European countries, the forestry related greenhouse gas inventories are largely or exclusively based on converting tree volume data from national forest inventories to biomass using biomass conversion and expansion factors (BCEFs). However, country specific data for many species are often lacking, which considerably increases the uncertainties of the greenhouse gas inventories. The focus of this research was to develop, using internationally published datasets that cover a large geographical area, an extended set of generalized curves of such biomass expansion factors for several species or species groups by age, growing stock and site index.     

Rootstock effects in grafted conifers: A review

The literature on rootstock effects (on scions) in conifers was reviewed, specifically: graft success, compatibility, size, reproduction, phenology, crown and needle characters, mineral contents, organic compounds, water relations, disease resistance and wood properties. Scions usually had higher graft success and less incompatibility on more closely related rootstocks although there were exceptions. Even intergeneric grafts have succeeded on occasion. Although there were marked rootstock effects on growth and reproduction, the effects did not follow a pattern with increasing relationship. It is also likely that some crown characters and the nutrient content of scions can be manipulated by the use of rootstocks. For many characters, a specific rootstock may give a desired result only for a limited number of scion types (species, cultivars or clones). With some exceptions, the review shows that the subject has not been comprehensively studied. Many of the studies were either short-term, inadequately replicated, or poorly designed to allow firm conclusions about rootstock effects. The physiological and biochemical mechanisms, which cause the changes seen in morphology, are not well understood. Further research and more comprehensive study of rootstock effects on scion biology are recommended.     

Ecology of Armillaria species on conifers in Japan

Distribution, host preference and pathogenicity of Japanese Armillaria species on conifers were investigated on the basis of field collections of 65 isolates. We identified seven Armillaria species from 19 conifer species including six major Japanese plantation conifers using mating tests and sequences of the translation elongation‐1 α gene. Armillaria mellea, Armillaria ostoyae, Armillaria cepistipes and Armillaria sinapina were frequently collected, whereas Armillaria nabsnona, Armillaria tabescens and a biological species Nagasawa’s E were rare. On the basis of host condition when the isolates were collected, A. mellea, A. ostoyae, A. cepistipes and A. tabescens are considered as moderate to aggressive pathogens of conifers in Japan.     

Tracheid length in relation to seedling height in conifers

Summary Changes in tracheid length during the development of primary and secondary xylem are analysed in relation to the division and elongation of cambial initials, in first year seedlings ofPinus sylvestris andPicea sitchensis. During primary and early secondary growth tracheid length varies with leaf internode length, but this relationship soon becomes obscured during later secondary growth. Correlations between seedling height and tracheid length thus depend on variation in tracheid length with radial xylem increment before and after terminal bud production. Since the relation between shoot and tracheid length is a complex one, the clonseness of the correlation varies with environmental treatment, but overall, a doubling of shoot length increased tracheid length by about 10%.I am grateful to ProfessorS. D. Richardson andDr. G. K. Elliott for many helpful suggestions. This work was financed by a grant from the Natural Environment Research Council. Plants were grown from seed kindly supplied by the Forestry Commission Research Division (Scotland and North England).     

Spatio-temporal variation of environmental signals inducing seed germination in temperate conifer plantations and natural hardwood forests in northern Japan

To document the spatial and temporal variation of environmental signals inducing seed germination in temperate forests, we measured temporal patterns of environmental signals and seed germination of six pioneer tree species in unthinned and thinned stands of conifer forests (Cryptomeria japonica plantations) and in the understory and gaps of hardwood forests in Japan. We also conducted germination experiment in laboratory for the six pioneer species to test the effects of red:far-red (R:FR) light ratio and temperature fluctuations on the seed germination. In conifer forests, the photosynthetic photon flux density (PPFD), the R:FR ratio, and the amplitude of temperature fluctuations in thinned stands were 2, 1.5, and 3 times higher, respectively, than those of unthinned stands. The PPFD and R:FR ratios just above forest floor also increased after the removal of thick litter accumulation. As a result, higher seed germination was observed in thinned compared to unthinned stands for three photoblastic species, whereas little differences were observed for three non-photoblastic species. These findings suggest that thinning, which frequently reduces litter accumulation, can substantially affect the regeneration of pioneer species and the resultant species diversity in conifer plantations. None of the measured environmental signals changed seasonally in unthinned stands of conifer forests, but they all changed remarkably in the understory of the hardwood forests. In this system, all signals were high and nearly identical to those in the gaps in early spring prior to canopy closure. Thus, the percent germination of the three photoblastic species was enhanced by high R:FR ratios and/or large temperature fluctuations even beneath the canopy and was nearly equal to that in the thinned conifer stands where the environmental conditions were nearly identical to those in the gaps. However, all of the environmental signals decreased with the expansion of canopy leaves and reached minimums at canopy closure. Even in the thinned stands and the gaps, the PPFD and magnitude of temperature fluctuations decreased over time due to shading by growing herbs and/or emerging canopy leaves. In these temporally changing environments, the germination of all photoblastic species ceased simultaneously. This study clearly demonstrated that the environmental signals inducing seed germination of photoblastic pioneer species spatially and temporally change in temperate forests, particularly in deciduous hardwood forests. Furthermore, these signals, PPFD, R:FR ratio, and the amplitude of temperature fluctuations, appear to play a very important role in tree regeneration and subsequent species diversity.