A diameter increment model for Crimean pine (Pinus nigra Arnold) and Calabrian pine (Pinus brutia Ten.) stands in Isparta region, Turkey

The purpose of this research was to test the precision of a diameter increment model for the estimation of future periodic diameter increment. Individual trees of Crimean pine (Pinus nigra Arnold) and Calabrian pine (Pinus brutia Ten.) located in both natural and plantation stands were selected. For that reason, normal closed canopy, pure, even-aged and undisturbed stands were examined. In 2002, plots were sampled in three natural and three plantation stands in Isparta region of Turkey. The number of sampling points in sample plots ranged from 19 to 55. In each sampling point, a subject tree and six competitors were selected. In each sampling point, subject tree and competitor trees were stem mapped (x and y coordinate system), and diameter (dbh), total height, age, and 10-yrs radial increment recorded. The predictors of a distance dependent diameter increment model were chosen that included tree level (diameter (d), competition index (CI), and age (t)) and stand level (basal area (G), and site index (SI)) characteristics as well as their transformations. The best fit index of the regression model was pursued in trials with variable combinations. The models explained 65%, 60%, 68% and 50% of the variation in individual tree diameter increment of Crimean pine and Calabrian pine for both natural and plantations stands, respectively. These models can be estimated diameter increment of individual trees at highly significant level (p<0.001).     

Genetic analysis of Pinus banksiana and Pinus resinosa populations from stressed sites contaminated with metals in Northern Ontario (Canada)

The Sudbury region in Canada is known for the mining and smelting of high-sulphide ores containing nickel, copper, iron and precious metals. Although reports provide information of metal levels in soil and plants, knowledge of genetic effects on plants growing in contaminated areas is limited. The main objective of this study was to characterise the level of genetic diversity in Pinus banksiana and Pinus resinosa populations from the Sudbury (Ontario) region using microsatellite markers. Soil samples were analysed for concentrations of metals. High levels of metal contents in soil were observed within short distances of the smelter compared with control sites. The level of genetic diversity was very low for P. resinosa populations and moderate for P. banksiana samples. Observed heterozygosity was fivefold higher in P. banksiana populations than P. resinosa populations studied. Overall, 17 and 24% of the total genetic diversity were attributed to differences among populations for P. banksiana and P. resinosa, respectively. In general, the inbreeding was significantly higher in P. resinosa populations than P. banksiana populations and gene flows were relatively low in both species. No significant trend of the levels of genetic diversity for metal contaminated and uncontaminated sites was found.     

Soil properties and root biomass responses to prescribed burning in young Corsican pine (Pinus nigra Arn.) stands

Fire is an important tool in the management of forest ecosystems. Although both prescribed and wildland fires are common in Turkey, few studies have addressed the influence of such disturbances on soil properties and root biomass dynamics. In this study, soil properties and root biomass responses to prescribed fire were investigated in 25-year-old corsican pine (Pinus nigra Arn.) stands in Kastamonu, Turkey. The stands were established by planting and were subjected to prescribed burning in July 2003. Soil respiration rates were determined every two months using soda-lime method over a two-year period. Fine (0-2 mm diameter) and small root (2-5 mm diameter) biomass were sampled approximately bimonthly using sequential coring method. Mean daily soil respiration ranged from 0.65 to 2.19 g Cm(-2) d(-1) among all sites. Soil respiration rates were significantly higher in burned sites than in controls. Soil respiration rates were correlated significantly with soil moisture and soil temperature. Fine root biomass was significantly lower in burned sites than in control sites. Mean fine root biomass values were 4940 kg ha(-1) for burned and 5450 kg ha(-1) for control sites. Soil pH was significantly higher in burned sites than in control sites in 15-35 cm soil depth. Soil organic matter content did not differ significantly between control and burned sites. Our results indicate that, depending on site conditions, fire could be used successfully as a tool in the management of forest stands in the study area.     

Effect of pine mistletoe on radial growth of Crimean pine (Pinus nigra) in Turkey

In this study, the influence of infection by pine mistletoe (Viscum album L. subsp. austriacum (Wiesb.) Volmann) on the radial growth of crimean pine (Pinus nigra Amold) in Turkey was investigated. We built local residual tree-ring-width chronologies using dendrochronogical techniques. Tree ring chronologies of uninfected (control) crimean pine were used to estimate potential radial growth characteristics in the "infected" crimean pine (light, moderate and severe infection groups). In 2005, increment cores were collected from 26 infected and 19 control dominant or co-dominant trees and annual radial growth indices from 1930-2005 were calculated for each infection group in a 14 point sampling. We compared radial growth in the uninfected trees with mean regional chronology. We found a strong decrease in radial growth in during the 1998-2005 period. The periodic average radial growth reduction (in %) from 1998 to 2005, respectively, were 0 for control, 26 for light, 39 for moderate and 63 for severe infection groups. It can be especially concluded that a severe degree of pine mistletoe attack has a negative effect on radial growth of the infected crimean pine trees.     

Removal of congo red and basic violet 1 by chir pine (Pinus roxburghii) sawdust,a saw mill waste: batch and column studies

The efficiency of chir pine sawdust (CPS) for adsorptive removal of the dyes, congo red (CR) and basic violet 1 (BV), from aqueous solution was evaluated using batch and column studies. The equilibrium was attained in 60 min for CR and 45 min for BV. The adsorption of BV obeyed the Langmuir isotherm model while the Freundlich isotherm fitted the equilibrium data of CR adsorption. The Langmuir monolayer adsorption capacities (Q_o) of CPS for BV and CR were 11.3 and 5.8 g kg^?1, respectively. The kinetic data for CR were best fitted to the Lagergren pseudo-first-order model and for BV to the pseudo-second-order model. The intra-particle diffusion was found to be the rate-controlling step for CR adsorption, while the adsorption kinetics of BV were controlled by both intra-particle and liquid-film diffusion. The thermodynamic parameters indicated that the adsorption process was spontaneous and endothermic in nature. The adsorption activation energy (E_a) for CR (124 kJ mol^?1) implied chemical adsorption while that for BV (5.4 kJ mol^?1) indicated physical adsorption. An increase in the Thomas model constant (K_Th) with increasing flow indicated that for both dyes the mass transport resistance decreased during adsorption. Thus, CPS may be an efficient low-cost adsorbent for decolorization of dye-containing wastewaters.     

Litter and dead wood dynamics in ponderosa pine forests along a 160-year chronosequence.

Disturbances such as fire play a key role in controlling ecosystem structure. In fire-prone forests, organic detritus comprises a large pool of carbon and can control the frequency and intensity of fire. The ponderosa pine forests of the Colorado Front Range, USA, where fire has been suppressed for a century, provide an ideal system for studying the long-term dynamics of detrital pools. Our objectives were (1) to quantify the long-term temporal dynamics of detrital pools; and (2) to determine to what extent present stand structure, topography, and soils constrain these dynamics. We collected data on downed dead wood, litter, duff (partially decomposed litter on the forest floor), stand structure, topographic position, and soils for 31 sites along a 160-year chronosequence. We developed a compartment model and parameterized it to describe the temporal trends in the detrital pools. We then developed four sets of statistical models, quantifying the hypothesized relationship between pool size and (1) stand structure, (2) topography, (3) soils variables, and (4) time since fire. We contrasted how much support each hypothesis had in the data using Akaike's Information Criterion (AIC). Time since fire explained 39-80% of the variability in dead wood of different size classes. Pool size increased to a peak as material killed by the fire fell, then decomposed rapidly to a minimum (61-85 years after fire for the different pools). It then increased, presumably as new detritus was produced by the regenerating stand. Litter was most strongly related to canopy cover (r2 = 77%), suggesting that litter fall, rather than decomposition, controls its dynamics. The temporal dynamics of duff were the hardest to predict. Detrital pool sizes were more strongly related to time since fire than to environmental variables. Woody debris peak-to-minimum time was 46-67 years, overlapping the range of historical fire return intervals (1 to > 100 years). Fires may therefore have burned under a wide range of fuel conditions, supporting the hypothesis that this region's fire regime was mixed severity.     

Effect of defoliation by the pine processionary moth (PPM) on radial, height and volume growth of Crimean pine (Pinus nigra) trees in Turkey

In this study, we assessed the effects of chronic defoliation on radial, height and volume growth of Crimean pine (Pinus nigra Arnold) trees of the pine processionary moth [Thaumetopoea wilkinsoni Tams (Lepidoptera:Thaumetopoeidae)] in western Turkey Crimean pine tree ring chronologies were analyzed for evidence of the pine processionary moth (PPM). Tree ring widths from non-defoliated Crimean pine sample trees, which were not defoliated by PPM from 1998 to 2004, were used to estimate potential growth in the defoliated Crimean pine sample trees during the same time interval. In 2004, increment cores collected from 50 defoliated sample trees and 25 non-defoliated sample trees dominant or co-dominant trees. Annual radial growth indices from 1985-2004 calculated for each defoliated Crimean pine and non-defoliated Crimean pine group. We identified regional outbreaks of PPM by synchronous and sustained growth periods of Crimean pine trees. Growth functions of defoliated Crimean pine trees (3) and non-defoliated Crimean pine trees (2) were graphically compared as the cumulative sum of radial, height and volume increment. Two outbreak were identified in 1992 (1992 and 1993) and 1998 (1998-2004) in the study area. PPM caused a significance decrease (average 33%, p<0.05) in the annual radial increment in 1998-2004.     

Whitebark pine (Pinus albicaulis) assisted migration potential: testing establishment north of the species range

The translocation of species into habitable locations outside of their current ranges, termed assisted migration, has been proposed as a means of saving vulnerable species from extinction as a result of climate change. We explore the use of this controversial technique using a threatened keystone species in western North America, whitebark pine (Pinus albicaulis), as a case study. Species distribution models predict that whitebark pine will be extirpated from most of its current range as temperatures rise over the next 70 years. However, the same models indicate that a large area within northwestern British Columbia, Canada, is climatically suitable for the species under current conditions and will remain so throughout the 21st century. To test the capacity of whitebark pine to establish relative to climatic and habitat features within its predicted climatic range, we planted seeds from seven populations in eight locations spanning from 600 km southeast to 800 km northwest of the northern boundary of the current species range. During the first three growing seasons, germination occurred in all locations. Nearly three times as many treated (induced maturation and broken dormancy) than untreated seeds germinated, and most treated seeds germinated a year earlier than the untreated seeds. Germination, survival, and growth were primarily influenced by seed mass, site climate conditions related to the duration of snow cover, and provenance temperature. Our experiment provides a preliminary test of models predicting the existence of climatically suitable whitebark pine habitat north of the current species ranges. More broadly, our techniques and results inform the development of scientific guidelines for assisting the migration of other species that are highly threatened by climate change. Applied case studies of this kind are critical for assessing the utility of species distribution models as conservation planning tools.     

山西高原油松种群遗传多样性

用酸性聚丙烯酰胺凝胶电泳（A-PAGE）技术，分析了山西高原9个油松种群在醇溶蛋白水平上的遗传多样性。135份材料共分离出23条带，其中3条为共有带，多态性高达86．95％。全部材料共出现53种带型，9个不同油松种群的带型有差异，同一种群不同个体的带型也有所不同，说明山西高原的油松在遗传上已产生一定程度分化，在醇溶蛋白水平上呈现出遗传多态性。从供试材料的带型计算出油松遗传分化系数为0．1547。即在种群间的变异占总变异的15．47％，种群内变异为84．53％，大部分的遗传变异存在于种群内，但种群间的分化程度在松属树种中也属于较高水平。根据23个多态位点计算遗传相似系数和遗传距离，进行聚类分析，将山西高原9个油松种群聚为3个类群。     

Dynamics of pheromone production and communication in the mountain pine beetle, Dendroctonus ponderosae Hopkins, and the pine engraver, Ips pini (Say) (Coleoptera: Scolytidae)

Summary. The mountain pine beetle, Dendroctonus ponderosae Hopkins, and the pine engraver, Ips pini (Say), often co-exist in lodgepole pine, Pinus contorta var. latifolia Engelmann. Intra- and interspecific semiochemical communication occurs in both species and their complete semiochemical repertoire and precise dynamics of pheromone production have not been elucidated. Porapak-Q extracts of captured volatiles from beetles of each species aerated at different attack phases (freshly emerged, pioneer sex alone in the log and both sexes paired in new galleries), followed by gas chromatographic-electroantennographic detection (GC-EAD) and GC-mass spectroscopic analyses identified 17 compounds (seven compounds common to both species, six present in D. ponderosae and four present in I. pini) that excited the antennae of either or both species. Seven compounds for D. ponderosae and nine for I. pini had not been assessed for behavioural activity. In field trapping experiments, 2-phenylethanol produced by both species inhibited the response of D. ponderosae to its aggregation pheromones. exo- and endo-Brevicomin produced by D. ponderosae significantly decreased the response of I. pini to its aggregation pheromone ipsdienol. Nonanal, a ubiquitous compound found in the volatiles of lodgepole pine, various nonhosts and in both beetle species deterred the response of I. pini to ipsdienol. The occurrence of cis-verbenol, trans-verbenol and verbenone in emergent I. pini, and verbenone and 2-phenylethanol in emergent D. ponderosae suggests that these compounds may inhibit aggregation and induce dispersal following emergence. Termination of aggregation in D. ponderosae appears to depend on the production of frontalin in combination with changes in the relative ratios of verbenone, exo-brevicomin, trans-verbenol and 2-phenylethanol. In I. pini, the cessation of ipsdienol production by males is probably the main factor in terminating aggregation. Received 16 November 1999; accepted 7 August 2000     

Carbon economy and nutrition of the alloparasitic red alga Harveyella mirabilis

Rhodomela confervoides (Huds.) Silva (Ceramiales: Rhodophyta) infected with the alloparasitic Harveyella mirabilis (Reinsch) Reinke (Cryptonemiales: Rhodophyta) was investigated with particular regard to rates of carbon fixation, patterns of soluble and insoluble assimilates, and transfer of reduced carbon from the photoautotrophic basiphyte to the fully heterotrophic epiphyte by means of ¹⁴C-labelling experiments. Gross carbon input by photosynthesis in Rh. confervoides accounted for about 53 g Cd^-1 when based on a biological unit (average individual of 200 mg fresh weight). Some 3% of the remaining net carbon gain (equivalent to about 0.7 g Cd^-1) was transferred from Rh. confervoides to its alloparasite, H. mirabilis (biological unit 0.5 mg fresh weight). Chromatographic examination showed that soluble ¹⁴C was translocated to and stored by H. mirabilis as mannosidoglycerate (=digeneaside) and probably a variety of amino acids. A certain proportion of soluble radiocarbon imported is converted by H. mirabilis to glycerolgalactoside (=floridoside). The bulk of imported ¹⁴C was recovered from the polymeric fraction, mainly from floridean starch.     

Possibilities for management of coastal foredunes with deteriorated stands ofAmmophila arenaria (marram grass)

Ammophila arenaria (marram grass) is the most important plant species for sand stabilization in European coastal foredunes. Stand degeneration due to poor supply of wind-blown sand enhances the susceptibility for wind erosion when successional species do not become established. ReplantedA. arenaria often fails to become established. In the present study we examined whether management practices such as mowing, fertilizing, burning or below-ground cutting of plants may be applied to re-establish the vigour ofA. arenaria. Field experiments performed at exposed sites, where naturally succeeding plant species are not supposed to become established due to salt spray, showed that none of the applied methods resulted into enhanced tiller density ofA. arenaria. Thus, further studies are necessary to solve this type of management problem. At the leeward side of foredunes, the successional speciesFestuca rubra ssp.arenaria andElymus athericus could be established successfully as both pre-grown seedlings and planted bundles of cuttings (all further experiments were fertilized). When planted in spring, cuttings of successional plant species established less than those planted in early winter. Water repellency of the sand surface is supposed to be the main cause for this. Pre-grown seedlings were less susceptible for the season of planting. Direct sowing was not effective. Rabbit browsing had to be omitted to obtain successful establishment. When living rhizomes of successional plant species were still present in the soil profile, fencing alone turned out to be effective to re-establish vegetation at initially bare sites.     

活性碳纤维的应用

本文在比较活性碳纤维与活性碳的结构和吸附特点的基础上,综述了这种吸附材料在废水处理,饮用水净化,空气净化,溶剂回收等各个领域的应用。     

Genetic protein variation in red mullet (Mullus barbatus) and striped red mullet (M. surmuletus) populations from the Mediterranean Sea

Starch-gel electrophoresis of allozymes was used to differentiate the two red mullet species (Mullus barbatus L. and M. surmuletus L.) in the Mediterranean Sea and, further, to investigate the genetic stock structure of M. barbatus in the eastern Mediterranean area. Twenty putative enzyme-coding loci were examined in eight M. barbatus samples caught in the Aegean and Ionian Seas (Greece) and in the Gulf of Lion (France), and two M. surmuletus samples caught in the Aegean and Gulf of Lion. A high degree of genetic polymorphism was found in both species. Species-specific electrophoretic patterns were found in PGI* and PGM*. Estimates of variance of allele frequencies among samples (F _ST) and ² analyses both revealed significant differences (P < 0.05) among the M. barbatus samples. Most of the genetic variation was among samples regardless of region. The mean value of Nei's genetic distance between the two species was 0.329. Genetic distance among M. barbatus samples was low (maximum Nei's D = 0.012), with the sample from Platania differing most from other M. barbatus samples. This is probably be due to founder effects existing at this area. These results suggest that allozyme analysis may provide important information on the genetic structure of the red mullet to ensure sustainable management of this species. Received: 7 May 1997 / Accepted: 13 October 1997     

燕山典型流域两种造林树种生态系统碳储量及固碳潜力研究

选择燕山典型流域6个林龄序列的小叶杨（Populus simonii）和5个林龄序列的山杏（Prunus sibirica）主要造林树种为研究对象,利用时间替代空间样地测量法量化退牧还林后生物量碳储量、凋落物碳储量和土壤碳储量及生态系统碳储量的变化规律,同时以各组成碳库-林龄序列中的最大碳储量之和作为生态系统饱和碳储量,以未退牧的天然草地生态系统碳储量作为初始植被类型的碳储量,分析总结了退牧还林对生态系统碳储量和碳循环的影响。结果表明,退牧还林后生态系统的生物量碳储量、凋落物碳储量基本随退牧年限的增加而增加,土壤碳储量随退牧年限的增加呈现先减小后增加的趋势。在没有人为干扰的情况下,9、15、18、22及29 a生小叶杨林的生态系统碳储量分别为7147.45、7461.67、7509.895、8468.375及8247.85 g·m^-2,9、15、18、22及26 a生山杏林的生态系统碳储量分别为6695.44、6700.82、8011.86、8001.92及7981.92 g·m^-2;9、15、18、22、29及36 a生小叶杨林的生态系统固碳潜力分别为757.08、1071.3、1119.53、2078.01、1857.48及1312.21 g·m^-2,9、15、18、22及26 a生山杏林的生态系统固碳潜力分别为310.45、1621.49、1611.55、1591.55及757.08 g·m^-2。长期来看,研究区退牧还林对提高生态系统碳汇能力是可观的、积极的。研究结果对提高造林对碳汇影响的估测能力提供数据支持,也为政府参与国际全球气候变化的谈判提供一个很好的案例研究和科学根据。     

Longleaf pine (Pinus palustris) and hardwood dynamics in a fire-maintained ecosystem: A simulation approach

Longleaf pine (Pinus palustris) savannas of the southeastern U.S. represent an archetype of a fire dependent ecosystem. They are known to have very short fire return intervals (∼1-3 years) that perpetuate understory plant diversity (up to 50 species m⁻²), support pine recruitment, and suppress fire sensitive hardwoods. Understanding the relationships that regulate longleaf and southern hardwoods is especially critical. With decreased fire frequency, insufficient intensity, or lack of underground competition, a woody mid-story rapidly develops, dominated by fire sensitive trees and shrubs that in-turn suppress more fire dependent species (including pine seedlings). This may occur in forest gaps, where pine-needle abundance is diminished, reducing fire spread potential. The interactions between longleaf pine, hardwoods, forest fuels, and fire frequency are complex and difficult to understand spatially. The objective of this study was to develop a spatially explicit longleaf pine-hardwood stochastic simulation model (LLM), incorporating tree demography, plant competition, and fuel and fire characteristics. Data from two longleaf pine study sites were used to develop and evaluate the model with the goal to incorporate simple site-specific calibration parameters for model versatility. Specific model components included pine seed masting, hardwood clonal sprouting, response to fire (re-sprouting, mortality), and tree density driven competition effects. LLM spatial outputs were consistent with observed forest gap dynamics associated with pine seedling establishment and hardwood encroachment. Changes in fire frequency (i.e., fire probability = 0.35-0.05) illustrated a shift in community structure from longleaf pine dominated to a hardwood dominated community. This approach to assessing model response may be useful in characterizing longleaf ecosystem resilience, especially at intermediate fire frequencies (e.g., 0.15) where the community may be sensitive to small changes in the fire regime. Height distributions and population densities were similar to in situ findings (field and LIDAR data) for both study sites. Height distributions output by the LLM illustrated fluctuations in population structure. The LLM was especially useful in determining knowledge gaps associated with fuel and fire heterogeneity, plant-plant interactions, population structure and its temporal fluctuations, and hardwood demography. This is the first known modeling work to simulate interactions between longleaf pine and hardwoods and provides a foundation for further studies on fire and forest management, especially in relation to ecological forestry practices, restoration, and site-specific applications.     

Movement and feeding activity of red sea urchins (Strongylocentrotus franciscanus) adjacent to a kelp forest

Movement and feeding were studied in a population of red sea urchins, Strongylocentrotus franciscanus (Agassiz, 1863), found within and immediately seaward of a kelp forest offshore from Santa Cruz, California, USA. Mean sea urchin movements varied from 7.5 cm/day inside the kelp forest to over 50 cm/day at 15 and 100 m outside the kelp forest. The percentage of sea urchins feeding decreased from 66% inside the kelp forest to 16 and 15% at 15 and 100 m outside the kelp forest. These data indicate that movement by these sea urchins is a response to a low food supply.     

Influence of growth on reproductive traits and its effect on fertility and gene diversity in a clonal seed orchard of scots pine, Pinus Sylvestris L

This study was carried out in a clonal seed orchard of scots pine (Pinus sylvestris L.), to determine the difference and interaction for reproductive and growth characters among clones and its impact on fertility variation and gene diversity Numbers of female and male strobili, and height and diameter at breast height were studied on six grafts chosen randomly in each of the 27 clones for the purpose. One-way analysis of variance revealed large differences in both reproductive and growth characters among clones. The differences were higher in growth characters than in reproductive traits. There was significant phenotypic correlation among growth and reproductive characters. So, growth characters had a greater effect on male and female fertility Estimates of total fertility variation (Sibling coefficient = 1.012), status number (26.8) and relative gene diversity (0.981) were computed. Fertility variation among clones was low, which caused a high relative population size (99% of census number). The positive phenotypic correlation between growth and reproductive characters showed that enhanced growth rate could be effective in improving fertility and gene diversity of seed orchard crop. The results of the study have implications in breeding and selection of plus tree and populations, establishment and thinning of seed orchards of the species.     

Carbon black-benzo(a)pyrene complex-induced apoptosis and autophagy blockage in rat alveolar macrophages

Abstract

Cooking of foods and the burning of biomass and fossil fuels in stoves are the main sources of cooking fumes, with carbon black and polycyclic aromatic hydrocarbons as main components. The toxicity of carbon black and polycyclic aromatic hydrocarbons has been well studied individually, but the combined toxicity is much less understood. Carbon black can adsorb benzo(a)pyrene to form a complex which displays an altered physical form which in this study has been constructed to simulate particles in the cooking fumes and explore the combined toxic effect on rat alveolar macrophages. The complex-induced cell apoptosis and blocked cell autophagy flux compared with both individually. The mechanism of toxicity may be by intracellular reactive oxygen species generation, impairing the mitochondrial membrane potential and activating apoptosis signaling pathways.     

Distribution of Carabid Beetles (Coleoptera, Carabidae) across a Boreal Forest–Clearcut Ecotone

Abstract: We studied the occurrence of carabid beetles (Coleoptera, Carabidae) in boreal forest fragments, their edges, and adjacent clearcuts in central Finland. Beetles were collected with pitfall traps along transects extending 60 m from the edge into clearcuts and 60 m into forest interior. Our main findings were that (1) species richness was significantly higher in the clearcut than in the forest fragments, (2) clearcuts hosted many open-habitat species, which increased overall species richness in these sites, (3) carabid assemblages in the edges were more similar to forest assemblages than to those found in the clearcuts, (4) no edge specialists were found, and (5) open-habitat species did not penetrate into the forest fragments from the clearcut. Because forest specialists occurred all the way to the edge on the forest side, it seems that edge effects per se do not adversely affect these species, at least in the short term. In the long term, however, habitat conditions in the edges may deteriorate for interior species because of trees falling over in strong winds, thereby reducing the size of the fragments and widening the edge zone.