Healing ability and diameter growth of lime-trees (<Emphasis Type="Italic">Tilia begonifolia</Emphasis> Stev.) following logging wounds

Lime-trees (Tilia begonifolia Stev.) have countless ecological and economical values in the northern forests of Iran. Bole wounds on residual trees are commonly caused by logging. The healing ability of bole wounds and the effect of bole wounds on diameter growth in lime-trees were assessed in a natural mixed uneven-age stand. The susceptibility to decay in relation to the wound severity was also evaluated. Condition and location of lime-trees before and after selective logging were determined through systematic plot sampling. The position of each lime-tree was identified on a topographical map using the global positioning system. After 10 years from wound occurrence, tree and wound characteristics were also re-measured. Overall, 68 stems (9.7 stem ha^?1) of lime-trees were found in the sampled area, 25 stems (36.8%) were damaged in the form of bole wound, and 43 stems (63.2%) were undamaged. Statistical differences between wounded or non-wounded lime-trees in terms of diameter and height were not found. Most of the wounds occurred at the height of < 1 m; the average wound size was 370.5 cm². Wound severity was related to wound size and wound position. Wound width healing rate was 6.2 mm year^?1. The wound shape was found to be a healing factor as the healing rate by width was higher than that by length. Healed wounds were only 12, 60% of bole wounds resulted in decay. Diameter growth of wounded trees was 43.5% lower than undamaged trees. The average annual diameter growth decreased by increasing wound severity. The results indicated the lime-tree is a very sensitive species to damage caused by logging (bole wound). Single-tree selection cutting needs more careful logging operations to reduce damage on residual lime-trees in order to safeguard this species, valuable in ecologic and economic terms.     

Observations on bark-stripping by red deer in a Picea sitchensis forest in Western Scotland over a 35-year period

The impact of bark-stripping by red deer on an 1825?ha plantation of Picea sitchensis is reported. Monitoring began in 1978 with initially 6500 trees at 37 sites; by 2014, only 10 sites remained due to felling. Sites became vulnerable to bark-stripping once the trees reached 7 years’ age, and from then until felling at age c. 45 years, incidence rates averaged 1% of the trees damaged yearly. Intense damage at a site in a year (>2% trees damaged) was often followed by intense damage next year, but, on average, 53% of sites were undamaged in a year. Many trees suffered repeat wounding, which reduced final impact by c. 30% compared to the theoretical impact calculated from annual damage and damage duration. The trees bark-stripped were on average smaller than plot mean girths once these means exceeded 20?cm; this had only a minor effect on the final impact since bark-stripped small trees had similar mortality to undamaged small trees. Most wounds (>90%) were small (<180?cm²) and healed quickly, so were unlikely to develop decay. Larger wounds never healed in less than 8 years, but the proportion healed increased progressively from 10 to 20 years after wounding.     

Discoloration following bark stripping wounds on Fraxinus excelsior

A total of 101 Fraxinus excelsior trees 8–32 cm in d.b.h. with bark stripping wounds caused by Alces alces and Cervus elaphus in Lithuania were cut, dissected and examined. The injuries ranged in age from 5 to 18 years, when the trees were 22–40 years old and were in the size range 6–24 cm d.b.h. For 5-10 years old wounds, the mean length of associated wood discoloration was 804 × 201 cm, while in 16–18 years old wounds it was 912 × 144 cm. There was no correlation between wound age and extent of discoloration in the stems examined (r = 0.11). Tree d.b.h. and wound size (initial width × length) were correlated with the extent of discoloration (r = 0.43 and r = 0.20, respectively). The presence of complete wound occlusion had no significant effect on the length of columns of discoloration associated with wounds of 10 or more years old. The average occlusion rate during the first 10 years was 128 × 28 mm or 13 mm/year, correlating positively with the radial increment (r = 0.42). Amongst a total of 500 F. excelsior trees with wounds, 29 trees (5.8%) bore basidiomycete fruiting bodies. Most common was Phellinus punctatus, found on 12 (2.4%) stems, followed by Coniophora puteana (1.6%), Peniophora limitata (0.4%) and Datronia mollis (0.4%). Fungi most commonly isolated from discoloured wood were ascomycetes Phaeoacremonium rubrigenum, Phialophora malorum, Hypoxylon rubiginosum, Coniothyrium sporulosum, Libertella spp., Ophiostoma sp., and basidiomycetes Ischnoderma benzoinum, Ischnoderma resinosum, Tyromyces sp.     

Treatment of wounds on Acer rubrum with Trichoderma viride1

Wounds on 90 Acer rubrum trees were inoculated with Trichoderma viride in July and August 1973 to determine the effect of the fungus on the development of decay. The trees were 45 years old and 15 cm diam at 1.4 m aboveground. Each tree had 12 wounds, 1.4 cm in diam and 5 cm deep. After a year 28 trees were harvested and isolations were made from the discolored, and decayed wood associated with each wound. Trichoderma viride was reisolated from all wounds that were inoculated with it. Hymenomycetes were isolated frequently from decayed wood associated with wounds not inoculated with T. viride. No Hymenomycetes were isolated from wounds inoculated with T. viride.     

Bark Stripping Damage by Red Deer in a Sitka Spruce Forest in Western Scotland: II. Wound Size and Position

Wounds to bark were measured within a few months of being inflictedby red deer in Glenbranter Forest, Argyll. About 8000 treeson 240 permanent plots were monitored; all the wounds occurringover a period of seven years were examined. Wound length varied little between forest stages, averaging12 cm. Wound breadth was greater on older trees, averaging 5cm in Sitka spruce and 6 cm in Norway spruce for the pole andhigh-canopy^* forest stages. A few wounds were large, maximumlengths being 95 and 80 cm in Sitka spruce and Norway sprucerespectively, but less than 20 per cent of the Sitka sprucewounds exceeded 100 cm² in surface area. Seasonal variationin size was modest; wounds were longest in spring (April–June). Often a single tree received two or more wounds during the 3-or6-month-long observation periods. For Sitka spruce there were89 instances of multiple wounding and 142 of single wounding,but these single wounds contributed only 37 per cent of thetotal of 383 wounds observed. The area of trunk wounded wasmore dependent on the size of the largest wound than on woundnumber, multiple wounds often being small. Trunks received most damage between 50 and 100 cm above ground,the lower and upper limits of wounding being 2 and 150 cm. Thepeaking in the incidence of damage with regard to height wasmost marked in pole-stage forest, whereas wounds were spreadmore evenly up the trunks on younger trees. Wound height waslittle affected by season, mean basal heights being 68, 70,74 and 67 cm in winter, spring, summer and autumn. Woundingon the trunk was not correlated with aspect.     

Wound occlusion and decay in Picea abies stems

In forest stands, Picea abies is commonly subjected to logging and bark stripping injuries. Most mechanical wounds do not exceed 300 cm², but their pathological consequences are unclear. The aim of the study was to estimate the rate of wound occlusion and the probability of decay in P. abies stems in relation to initial wound size and wounding season. The study included forty-five stems with 1-, 2-, 3-, 4- and 5-cm-wide by 15-cm-long wounds (15–75 cm²), and eighty-three stems with 15-cm-wide by 20-cm-long wounds (300 cm²), made either in January or in August. After 19–21 years, each wound was assessed for occlusion rate and the presence of decay. All 1- to 2-cm-wide wounds were occluded, compared to 58–70% of the 3- to 4-cm-wide wounds, 25% of the 5-cm-wide wounds, and 0% of the 15-cm-wide wounds. On average, it took 3.6, 5.5, 10.4, 12.7 and 14.7 years to occlude wounds ranging in size from 1- to 5-cm-wide, respectively. After 21 years, the observed size of wounds comprising an initial area of 300 cm² ranged between 14 and 481 cm². Wounding season and stem DBH had no impact on wound healing. Wound decay incidence correlated positively with the initial width of injury (r = 0.925; P < 0.05). All 1- to 2-cm-wide wounds lacked any decay, while the proportions of decay among 3-, 4-, 5- and 15-cm-wide wounds were 58.3, 50.0, 83.3 and 100%, respectively. Length of decay comprised 35–225 cm. Consequently, wounds on P. abies stems greater than 5 cm width are unlikely to be occluded and more prone to decay development.     

BACTERIAL SWOLLEN STEM CANKER ON POPLAR:THE DISEASE AND ITS CAUSES

The bacterial swollen stem canker of poplar is prevalent in planted poplars ofnortheast China.Serious damage occurs in the form of necrosis,putridity,cankers,swollen stemsand discolored wood of trunks and branches of young and mature trees.Tests of allergic reaction,pathogenicity,culture characteristic,Gram stain and anaerobic culture revealed that the pathogen isErwinia sp.Its occurrence depends mainly on the predisposing factors of the disease among whichthe susceptibility of poplars is the most important one.The disease becomes widespread in associa-tion with low-lying and high-humidity terrain,frost injury,node wounds,insect damage,and prun-ing and other mechanical wounds.Harmful factors promote the disease.     

The characteristics of oleoresin exudation amount from the trunks of pine trees

The amounts of oleoresin exudation from artificially-marked wounds on the trunks of pine trees were measured, and several characteristics of the oleoresin exudation amount (OEA) were elucidated. OEA showed no regular diurnal change and had no relation to both heights and directions of the wounds marked on the trunks, but differed between individual trees and between locations on the same tree. The exudation of oleoresin from each wound stopped after several hours. Mean OEA per wound decreased with increase in the number of wounds per given area. When the exudation of oleoresin from a wound was exhausted, further oleoresin was scarcely exuded even from a new wound made close to the initial one. In response to wounding, therefore, primary oleoresin exudation seems to occur by releasing the resin stocked in the resin canal.     

An investigation of conditions occurring within pruning wounds

The electrical resistance at various points within pruning wounds was measured, over periods of up to two years, using a pulsed electric current resistance meter (Shigometer). Wood with stain and decay generally had a lower electrical resistance than healthy wood. Mathematical and visual models were constructed from measurements of electrical resistance and examination of dissected wounds. These models were used to relate the extent of stain and/or decay with the diameter of the wound, length of the wound stub, time after pruning and with wound treatment.     

How fast do European conifers overgrow wounds inflicted by rockfall?

The capacity of trees to recover from mechanical disturbance is of crucial importance for tree survival but has been primarily investigated in saplings using artificially induced wounds. In this study, mature Larix decidua Mill., Picea abies (L.) Karst. and Abies alba Mill. trees growing on alpine slopes that were wounded by naturally occurring rockfall were analyzed to determine their efficiency in overgrowing wounds. In total 43 L. decidua, P. abies and A. alba trees were sampled. First, 106 samples from 27 L. decidua and P. abies trees were analyzed to reconstruct yearly and overall overgrowth rates. Cross sections were taken at the maximum extension of the injury and overgrowth rates were determined on a yearly basis. Results clearly showed that L. decidua overgrew wounds more efficiently than P. abies with an average overgrowth rate of 19° and 11.8° per year, respectively. The higher on the stem the injury was located, the faster the wound was closed. Young and small trees overgrew wounds more efficiently than older or thicker trees. In contrast, no correlation was observed between injury size or increment before/after wounding and wound closure. Second, cross sections from 16 L. decidua, P. abies and A. alba (54 injuries) were used to assess closure rates at different heights around the injury. Overgrowth was generally smallest at the height of the maximum lateral extension of the injury and increased at the upper and lower end of the injury. The efficiency with which L. decidua closes wounds inflicted by rockfall makes this species highly adapted to sites with this type of mechanical disturbance.     

The Development of Stain in Wounded Sitka Spruce Stems

In order to investigate the severity of staining in woundedSitka spruce stems, the vertical extent of two categories ofstain (based on colour and termed ‘light’ and ‘heavy’)was measured in stems at two sites in south Scotland. Data wereobtained from 98 wounds most of which were between 8 and 14years old and most of which had been caused by Red deer. Theupward extent of stain and its rate of upward spread were positivelycorrelated with wound surface area and length. Most stain wasof the light type which, although it was associated with nearlyall wounds, usually constituted an insignificant defect anddid not commonly extend for more than 1 m ahve wounds. Heavystain occurred above less than half of all wounds studied butwas common above wounds exceeding 300 cm² surface area. Forall wounds, the mean value for the upward extent of heavy stainwas 16.1 cm with a mean rate of spread of 1.9 cm y^–1.However, for wounds exceeding 300 cm² surface area, the valueswere 51.7 cm and 5.5 cm y^–1. These results suggest that,although there is an important effect of wound size on the typeand extent of stain, wounds of the type studied are unlikelyto lead to severe stain and decay in Sitka spruce over a periodof 8–14 years. The results are discussed in relation toother studies on wound staining in conifers.     

Impact of bark wounds on sapwood in Norway spruce and silver fir

Bark wounds by damage during harvesting are a serious problem in forestry due to fungi infection and wood deterioration. This paper presents results of an investigation about the influence of the wounds on the internal structure of such injured stems. In an experiment, bark wounds were artificially created at the stem base of Norway spruce [Picea abies (L.) Karst.] and silver fir (Abies alba Mill.). 2 years later, the injured stems along with undamaged controls were cut and the trunk portion below breast height subjected to computer tomographic (CT) analysis. Analysis of the CT-images revealed a substantial impact of wounding on sapwood properties in spruce: directly adjacent to the wound surface in all examined trees, a large disturbance zone was detected affecting on average 17% of the potential sapwood area. With increasing distance from the wound, the size of this disturbance zone diminished, but was still detectable in all trees at breast height ca. 1 m above the bark wound.     

Localized within‐ and between‐tree variation in nematode distribution during latent state of pine wilt disease makes the disease status cryptic

The pinewood nematode (PWN), Bursaphelenchus xylophilus, is the causal agent of pine wilt disease, an epidemic disease that has severely damaged pine forests in East Asia. The disease has spread to northern areas in Asia and parts of Europe. To prevent disease spread as the forefront of damage prevention, a better understanding of infection status is highly important. Not all infected trees show disease symptoms, and such asymptomatic PWN‐carrying trees are likely to be overlooked and can become a pathogen reservoir. To elucidate PWN infection status in asymptomatic trees, we performed PWN inspection of branches and trunks in 21 test trees in two different conditions: trees that had experienced PWN inoculation and those with suspected PWN infection that had experienced transient foliage discoloration. We detected PWNs in eight test trees (38%) and in 13 (1.5%) of a total of 843 samples. The difference in these percentages suggests that nematode inhabitation was highly localized within the trees, possibly owing to the restricted migration of PWNs. Our data demonstrated that trees that were once weakened but recovered their vigour can persist, as the asymptomatic carriers, in the forest. The implications for disease control are also discussed.     

Growth effects of thinning damage in a Corsican pine (Pinus laricio Poiret) stand in central Italy

Damage to residual stand after partial harvesting or thinning may lead to serious economic losses in terms of both timber quality at the final harvest, and tree growth reduction. Logging damages and their effect on tree growth were studied in a long term experiment on Corsican pine in central Italy. Damage frequency, agent (felling, skidding), position (root damage, stem base, between 0.3 and 1 m a.g.l., >1 m a.g.l.) and severity (light, severe) and tree growth were measured after selective thinning from below and at 10 years after the treatment. In detail, we aimed at: monitoring mechanical damages to trees at the end of thinning and after 10 years; and assessing stand stability, growing stock, ring width and basal area at 10 years after the thinning. The thinning removed about 20% of volume, 38% of trees and 26% of basal area. The basal area decreased from 56 m² ha⁻¹ to 42 m² ha⁻¹ but after 10 years it increased again to 56 m² ha⁻¹. Immediately after thinning, 13.6% of the standing trees was damaged, out of these 36.17% showed severe injuries. Damages to standing trees were mainly due to skidding. Ten years after thinning, the percentage of damaged trees was about 17%, out of which 86.67% showed severe wounds. An increase of damaged trees and of trees with severe wounds was observed suggesting that a deeper knowledge on long-term effect of logging damages is needed. This study did not highlight any effect of logging damage on tree growth. In fact, no difference in ring width was recorded between damaged and undamaged trees.     

Micromorphology of abnormal and decayed xylem in rubberwood canker

Typical symptoms of rubberwood canker found in East Kalimantan, Indonesia, differ from those of conventional rubberwood peach canker. Rubberwood canker showed a conspicuous longitudinal canker with exposed xylem that was discolored and decayed. Bark wounds, resulting from regular tapping, were enclosed within the xylem, and the tissues had become integrated. The characteristics of abnormal and decayed xylem in this rubberwood canker were analyzed by light microscopy and scanning electron microscopy. Xylem in the vicinity of the enclosed bark contained fewer vessels of smaller diameter and shorter length, and significantly wider rays compared with normal xylem. Around the wide growth zones of the canker, axial cells were disoriented and warped toward the canker zones. In view of the separation among cells, and the concentric degradation of the cell walls starting from the lumen surfaces, decayed xylem appeared to be caused mainly by white-rot fungal attack.     

Effectiveness of a detached‐leaf assay as a proxy for stem inoculations in backcrossed chestnut (Castanea) blight resistance breeding populations

A recently developed detached‐leaf blight resistance assay has generated interest because it could reduce the amount of time needed to evaluate backcrossed hybrid trees in the American chestnut blight resistance breeding programme. We evaluated the leaf inoculation technique on a sample of advanced progeny from the Indiana state chapter American Chestnut Foundation breeding programme, along with susceptible American chestnut (Castanea dentata), the recurrent parent, and resistant Chinese chestnut (Castanea mollissima), the donor parent for blight resistance. In experiments over 2 years using two pathogen isolates, we found no biologically meaningful relationship between leaf lesion size and the size (length and width) or severity (1–5 canker severity rating) of stem cankers on 5‐year‐old trees. Chinese chestnuts did develop significantly smaller leaf lesions than American or backcrossed chestnuts. We conclude that while the detached‐leaf assay may have utility in some chestnut breeding applications, it is not a suitable proxy for the established practice of stem inoculations.     

Responses ofQuercus sapwood to infection with the pathogenic fungus of a new wilt disease vectored by the ambrosia beetlePlatypus quercivorus

Quercus serrata andQ. crispula wilt during the summer in wide areas along the Sea of Japan. Mass attacks of trees by an ambrosia beetle (Platypus quercivorus) are characteristic before appearance of the wilting symptoms. This study investigated the pathogenic effects of a fungus detected specifically in the wilting trees. This hyphomycete fungus,Raffaelea sp., has a distribution that correlates with the discolored xylem area called wound heartwood in which vessels are dysfunctional. Tylosis formation around the hyphae indicates vessel dysfunction. In areas with discoloration, the fungal hyphae were invading living ray parenchyma cells from the vessel lumen. As a protective reaction the ray cells exuded yellow substances into the vessels, but these substances seemed ineffective against the fungal activity, probably because the fungus disperses along the beetle's gallery before enough substance can accumulate. It should allow wide discoloration in sapwood. Cambium was not necrotic around the fungus. The cytological process in the host was as follows: (1) synthesis of secondary metabolites by the stimuli of oak fungus; (2) exudation of yellow substances into vessels; and (3) dysfunction of vessels and wound heartwood formation. In regard to wilting of trees, the pathogenicity of the fungus should be assessed by its ability to stop sap flow.     

Bark phenolics of American beech (Fagus grandifolia) in relation to the beech bark disease1

The amount of total extractable phenols was determined for bark sections obtained from behind cankers naturally induced by Nectria coccinea var. faginata and from behind mechanically inflicted wounds on stems of American beech. Healthy bark from susceptible trees was found to contain a similar level of phenolics as bark from trees determined to be resistant to the beech bark disease. Six months after wounding, wound-altered bark from susceptible trees was found to be lower in phenols than woundaltered bark from resistant trees. Inoculation of wounds with N. coccinea var. faginata resulted in decreased phenolic levels in bark sections nearest the wound surface, and increased phenolic levels in sections nearest the vascular cambium, several mm distant. Phenolic levels in injured or infected bark appear to follow similar patterns as those resulting from injury or infection of xylem tissues.     

Seasonal variations in the incidence of pine wilt and infestation by its vector, Monochamus alternatus, near the northern limit of the disease in Japan

During the last few decades, pine wilt disease has spread to cool-climate regions in Japan and, more recently, the potential risk of it spreading into the European midwest has also become a concern. In a coastal pine stand (84.7?ha) in Akita, near the northern limit of pine wilt disease in Japan, we investigated seasonal variations in the incidence of damage caused by the disease to trees and oviposition by the disease’s insect vector, Monochamus alternatus, during a two-year period. Foliage discoloration occurred throughout each year, and its seasonal variation showed a bimodal pattern in Pinus thunbergii (a higher peak in May–June and a smaller peak in October) and a clear peak in June in P. densiflora, which differed from the patterns in seasonal variation seen for warm-climate regions. Oviposition scars by M. alternatus were found in 40–45% of the trees damaged each year. The percentage of trees that had oviposition scars was higher in P. thunbergii than in P. densiflora. This appeared to reflect the difference in seasonal discoloration pattern between the two species. Analysis of the oviposition risk showed that trees that exhibited discoloration starting between July and October had a significantly higher risk or significantly higher oviposition scar densities, particularly for those that became discolored between August and September (2.5–14.6-fold higher risk than during other months). Oviposition scar densities per damaged tree were similar within the period of higher oviposition risk. Considering both oviposition risks and scar densities, we concluded that trees with discoloration that become apparent between July and October are important targets for preventing the spread of pine wilt disease in Akita.     

Discoloration and associated fungi in stems of silver birch (Betula pendula Roth.) following logging damage

Study aim was to investigate discoloration and associated fungi in 50‐ to 70‐year‐old stems of Betula pendula damaged by logging. Investigated wounds (on 70 stems) were inflicted 4–18 years previously, and all of them had discoloration with mean radial spread (±SD) 2.9 ± 2.9 cm and longitudinal spread 50.3 ± 24.0 cm. Both wound area and length had a strong impact on a total length of discoloration (r = 0.665 and r = 0.745, respectively; p < 0.001), but had no influence on its spread beyond wound margins (r = ?0.035 and r = ?0.079). There was no correlation between the age of an injury and spread of the discoloration neither in vertical (r = ?0.110) nor in radial (r = ?0.280) direction. Average wound closure rate was 1.2 ± 0.5 cm year ^?1. The difference between discoloration length in stems with open and occluded injuries was insignificant (t‐test, p = 0.769). The principal fungi isolated from the wounds were ascomycetes (Epicoccum, Cadophora, Neonectria, Alternaria spp.) that were able to cause discoloration of wood with little effect on its mechanical properties and capable only for limited spread inside a stem of a living tree. In conclusion, results of the present study demonstrate that the amount of discoloured wood beyond logging wounds in B. pendula is mainly restricted to the damaged portion of a stem and that the associated loss of wood production remains low during about 10 years since the damage.