Lens cell-to-cell channel protein: I. Self-assembly into liposomes and permeability regulation by calmodulin

Summary Lens fibers are coupled by communicating junctions which contain a 28-kDalton protein (MIP26) believed to be the main component of the cell-to-cell channel. To study the permeability properties and regulation of these channels, anin vitro system has been developed in which MIP26 isolated from calf lens is incorporated into liposomes and the resulting channels are studied spectrophotometrically by a swelling assay. Liposome vesicles were prepared using a sonication/resuspension method. Incorporation efficiency was monitored by freeze-fracture. Vesicles were resuspended in 6% Dextran T-10. Assay buffer was identical, except for isotonic substitution of sucrose for T-10. MIP26-incorporated (but not control) vesicles swell under isotonic conditions indicating sucrose entry (via channels) followed by water to maintain osmotic balance. In the absence of calmodulin, calcium ion has no effect on channel permeability. On the contrary, vesicles prepared with equimolar amounts of MIP26 and CaM do not swell in the presence of calcium ion, indicating that the channels can be closed. Addition of EGTA to these vesicles reinitiates swelling—evidence that the channel gating mechanism is reversible. Magnesium ion has no effect on either type of vesicle.     

Backwater habitats and their role in nature conservation on navigable waterways

In Britain, lightly trafficked canals frequently contain diverse, productive macrophyte communities. These represent important habitats for macroinvertebrates and fish while having a high intrinsic value in nature conservation terms. As recreational boat traffic increases, fragile macrophytes are progressively eliminated and the biomass of the remaining species is greatly reduced, thereby adversely affecting weed-associated animals and ultimately simplifying the structure of the whole ecosystem. From the viewpoint of aesthetics, nature conservation and fisheries management,ecological enhancement of these traffic impacted ecosystems is desirable but options are limited by channel size and the intensity and type of disturbance. Backwater areas connected to the main channel but apparently remote from traffic influences ought however, to provide a minimally-disturbed refuge for macrophytes and dependent organisms. An extensive field survey was undertaken to test this hypothesis and evaluate the potential for exploiting backwater sites as off-line nature reserves. Principal determinants of vegetation structure and species diversity are identified and discussed and are used to prescribe a set of ideal characteristics for prospective backwater nature reserves and to forecast likely management problems.     

Resuspension in a shallow eutrophic lake

The frequency and the importance of wind-induced resuspension were studied in the shallow, eutrophic Lake Arresø, Denmark (41 km², mean depth 3 m). During storm events in autumn 1988 lake water samples were collected every 2–8 hours by an automatic sampler at a mid-lake station. The concentration of suspended solids and Tot-P was found to increase markedly. During storms up to 2 cm of the superficial sediment was resuspended, and the concentration of resuspended solids in the water column rose to 140 mg l^–1. The resuspended particles had a relatively high settling velocity and on average, a relatively short residence time in the water column of 7 hours.A model which describes the concentration of resuspended solids as a function of wind velocity and of settling velocity of the resuspended particles is presented. Using additional wind velocity data from a nearby meteorological station, the model has been used to calculate the frequency of resuspension events and concentration of resuspended solids for the period from May to November 1988.These calculations show that resuspension occurred about 50% of the time. Average flux of suspended solids from the sediment to the water was 300 g m^–2 d^–1 and during 50% of the time lake water concentration of suspended solids was more than 32 mg l^–1. A relationship between concentration of suspended solids and Secchi-depth is presented. Because of resuspension, Secchi-depth in Lake Arresø is reduced to 0.5 m.Resuspension also had a marked effect on Tot-P concentration in the lake water, and P input to the lake water being totally dominated by resuspension events.     

Empirical evidence of the importance of sediment resuspension in lakes

This paper reviews the empirical evidence for bottom sediment resuspension, which has been documented in many aquatic systems, and attempts to put those observations into a predictive framework. The evidence suggests that resuspension is episodic in all systems with the spatial extent of resuspension being determined by the wind speed, duration of disturbance, effective lake fetch and lake morphometry. The rates of deposition of resuspended material vary greatly with measurements recorded between 0.5 and 50 g m^?2 d^?1. In many systems and at many times of year resuspended material comprises the majority of the flux of particles to lake bottoms. An order-of-magnitude estimate of the long-term contribution of resuspended material to the total flux of particulate matter is 85%.     

Behavioural Effects of Tourism on Oceanic Common Dolphins,Delphinus sp., in New Zealand: The Effects of Markov Analysis Variations and Current Tour Operator Compliance with Regulations

Common dolphins, Delphinus sp., are one of the marine mammal species tourism operations in New Zealand focus on. While effects of cetacean-watching activities have previously been examined in coastal regions in New Zealand, this study is the first to investigate effects of commercial tourism and recreational vessels on common dolphins in an open oceanic habitat. Observations from both an independent research vessel and aboard commercial tour vessels operating off the central and east coast Bay of Plenty, North Island, New Zealand were used to assess dolphin behaviour and record the level of compliance by permitted commercial tour operators and private recreational vessels with New Zealand regulations. Dolphin behaviour was assessed using two different approaches to Markov chain analysis in order to examine variation of responses of dolphins to vessels. Results showed that, regardless of the variance in Markov methods, dolphin foraging behaviour was significantly altered by boat interactions. Dolphins spent less time foraging during interactions and took significantly longer to return to foraging once disrupted by vessel presence. This research raises concerns about the potential disruption to feeding, a biologically critical behaviour. This may be particularly important in an open oceanic habitat, where prey resources are typically widely dispersed and unpredictable in abundance. Furthermore, because tourism in this region focuses on common dolphins transiting between adjacent coastal locations, the potential for cumulative effects could exacerbate the local effects demonstrated in this study. While the overall level of compliance by commercial operators was relatively high, non-compliance to the regulations was observed with time restriction, number or speed of vessels interacting with dolphins not being respected. Additionally, prohibited swimming with calves did occur. The effects shown in this study should be carefully considered within conservation management plans, in order to reduce the risk of detrimental effects on common dolphins within the region.     

植物叶表附属物对大气颗粒物滞留及再悬浮的影响研究:以空气凤梨为例

植物叶表附属物是与大气颗粒物最先接触的结构,但现在少有研究表明这些特殊结构是如何影响大气颗粒物的滞留的。因此,本研究以具有典型叶表面附属物-鳞片的植物-空气凤梨维路提拉为实验对象,通过人工去除鳞片,比较鳞片的有无对其叶片滞尘量的影响,并比较了不同时间、不同风力强度条件下其叶片表面所滞留的大气颗粒物再悬浮的比例。结果表明,未去除叶表鳞片的维路提拉的最大滞尘量（23.24±0.11 g·m^-2）和自然滞尘量（10.22±0.92 g·m^-2）均显著高于去除鳞片的植株（P<0.05）。大气颗粒物沉降到叶片表面后,在风力作用下,去除鳞片的空气凤梨植株上超过99.0%的粉尘会被吹起,而未去除鳞片的空气凤梨植株上这一比例仅有28.1%。而且,鳞片去除的越多,大气颗粒物的再悬浮比例越大。以上结果表明维路提拉叶表鳞片对大气颗粒物的作用不仅体现在总滞尘量上,而且体现在大气颗粒物的再悬浮过程中,它会有效降低沉降于叶表面颗粒物的再悬浮,起着促进颗粒物滞留的作用。     

通道大小对经皮肾镜取石患者出血的影响

目的:探讨通道大小对经皮肾镜取石患者围手术期出血的影响及治疗策略。方法:回顾性分析2016年1月至2017年1月在我院行F24和F18通道的经皮肾镜取石术的189例患者临床资料,分别比较具有不同临床特征患者围手术期出血的发生情况。结果:95例F24通道患者平均出血量为125±19.6 m L,其中5人进行输血治疗;94例F18通道患者平均出血量为103±17.6m L,其中3人进行输血治疗。F18通道经皮肾镜取石术的出血组和非出血组糖尿病、高血压的发生率、结石表面积、通道数量、手术时间比较差异均具有统计学意义(P0.05)。F24通道经皮肾镜取石术的出血组和非出血组孤立肾、高血压发生率、结石面积、肾实质厚度、通道数量和手术时间比较差异均具有统计学意义(P0.05)。F18通道经皮肾镜取石围手术期出血量显著少于F24通道(P0.05)。结论:孤立肾、高血压、结石面积大、肾实质厚、肾积水轻、通道数量多和手术时间长均会导致经皮肾镜围手术期出血几率和出血量增加,并且F24通道相较于F18通道出血量更多。     

Short-term variations of seston characteristics in a shallow tropical lagoon: Effect of wind-induced resuspension

In January 1993, a 24 hour survey based on an hourly sampling was conducted in a station (1 m depth) located in the Ebrié lagoon, Côte d'lvoire, in order to determine short-term changes in seston quality and quantity and relate them to physical conditions. Water samples were collected just below the surface and at 0.9m depth to assess the seston distribution in the water column. Located in an estuarine environment, the study site was submitted to periodic effect of tides with low currents. Under certain conditions, sediment resuspension by wind-induced waves was observed in the shallow parts of this ecosystem. From the analysis of hydrodynamic processes (tide, wind and sedimentation rates), four basic situations were distinguished in the course of the study: a period with no wind and high sedimentation rates, a period characterized by wind-induced resuspension, followed by a period of sedimentation and finallly, a period without wind during the second flood tide. The particle size spectra were similar throughout the study, the distributions being unimodal (mode of 4 μm diameter). The finer particles (<10 μm) were resuspended over the whole water column at the beginning of the resuspension period, whereas the larger particles were resuspended if the wind speed exceeded 3 m s^?1. Phytoplankton species were found to provide information on the physical factors controlling the pelagic system: small cells, such as those of cyanobacteria (Synechococcus sp.), were always present over the whole tidal cycles, whereas large motile cells were predominant in the water column during the resuspension period. The seston characteristics of this part of Ebrié lagoon are mainly controlled by tidal currents, but wind-induced resuspension can modify the structure of the pelagic food web.     

Measurement of resuspended aerosol in the Chernobyl area

During anthropogenic activities, such as agricultural soil management and traffic on unpaved roads, size distribution measurements were performed of atmospheric particulate radionuclides at a site in the Chernobyl 30-km exclusion zone. Analysis of cascade impactor measurements showed an increase of the total atmospheric radioactivity. In the cases of harrowing by a tractor and traffic on unpaved roads, a common shape of the size distribution was found with two maxima, the first in the 2–4 μm range, the second in the 12–20 μm range. The size distributions were compared to measurements during wind-driven resuspension. Particle number concentration measurements with an Aerodynamic Particle Sizer showed a dynamic dependence of the particle concentration in different size ranges on anthropogenic action. The increase of the mean concentration was for the large particles more than one order of magnitude higher than for fine particles during anthropogenic enhanced resuspension. From the measurement of the mass concentration, the radioactive loading could be estimated. An enrichment of radionuclides on resuspended particles (compared to soil particles) was found, with the highest enrichment for large particles. Micrometeorological considerations showed that large particles may frequently be subject to medium range transport. The dry deposition velocity was measured; the mean value of 0.026 m s^–1±0.016 m s^–1 is typical for 6–9 μm diameter particles. Received: 17 February 1998 / Accepted in revised form: 17 June 1998     

Vessel collision injuries on live humpback whales,Megaptera novaeangliae,in the southern Gulf of Maine

North Atlantic humpback whales (Megaptera novaeangliae) in the Gulf of Maine overlap with both recreational and commercial vessel activity. Vessel strikes are one source of anthropogenic impact that has the potential to inhibit the recovery of this protected species. There are currently no regulations or guidelines specifically devised to reduce the likelihood of collisions for vessels transiting in the vicinity of humpback whales, except for vessels actively engaged in whale watching. To understand interactions between vessels and humpback whales better, we analyzed injuries on 624 individuals photographed in the southern Gulf of Maine from 2004 to 2013. Multiple reviewers evaluated 210,733 photos for five categories of injury consistent with a vessel strike. In total, 14.7% (n = 92) of individuals photographed showed injuries consistent with one or more vessel strikes. These results likely underestimate vessel collision rates and impacts because multiple events, events resulting in mortality, and those that involved only blunt force trauma could not be detected. Nevertheless, our results indicate that vessel strikes are underreported and that healing is dependent on the severity and location of the injury. We recommend that a management strategy be developed for all classes of vessels transiting in the vicinity of whales.     

Survey augmentation using commercial vessels in the Mid-Atlantic Bight: Sampling density and relative catchability

A series of side‐by‐side tows was conducted between a survey vessel and a commercial vessel in two seasons, spring and fall (autumn), to examine the use of commercial vessels to increase sampling density in trawl‐based stock surveys. Both vessels caught more fish offshore in the spring. The commercial vessel caught more fish than the survey vessel in both seasons. Catches of nearly all species were contagiously distributed in the spring. Most were contagiously distributed in the fall; however, somewhat more species were characterized by random or even distributions. The variance‐to‐mean ratio was consistently higher for most species for commercial vessel catches, regardless of season. As both vessels sampled in the same region at the same time, the increased predilection for the survey vessel to assess the distribution pattern as less patchy than the commercial vessel must accrue from some difference in sampling dynamics rather than variation in species distribution. A simulated decrease in sampling effort from 59 to 30 or 15 hauls increased the variance‐to‐mean ratio. Reduced sampling effort increased the tendency for occasional large catches to vary the estimate of domain biomass. The sampling program included an onshore–offshore gradient in station density. Domain biomass was considerably underestimated with reduced station density for six species characterized by large catches offshore in that portion of the survey domain characterized by low station density. In this study, a factor of two variation in domain biomass became more likely in 40% of species when sampling effort was reduced to 15 hauls from 59. A factor of two in biomass may distinguish a sustainable fishery from one in which a species is overfished. As survey sampling effort in this area was 18 hauls, increasing sample number by inclusion of commercial vessel tows would be advantageous. A regression between paired tows failed to adequately predict catches of one vessel from catches of the other. Standardization of vessel catches by the ratio‐of‐mean catches provided a more realistic comparison because large catches accounted for a significant fraction of domain biomass; however, a single conversion coefficient between boats could not be used for both sampling periods. The underlying impediment in developing a general conversion factor between the two vessels seems to be rooted in the differential in variance‐to‐mean ratios of the catches; this differential exists despite sampling of the same distribution of fish.     

Effects of reagents modifying carboxyl groups on the gating current of the myelinated nerve fiber

Summary K⁺ channels in inside-out patches from hamster insulin tumor (HIT) cells were studied using the patch-clamp technique. HIT cells provide a convenient system for the study of ion channels and insulin secretion. They are easy to culture, form gigaohm seals readily and secrete insulin in response to glucose. The properties of the cells changed with the passage number. For cell passage numbers 48 to 56, five different K⁺-selective channels ranging from 15 to 211 pS in symmetrical 140mm KCl solutions were distinguished. The channels were characterized by the following features: a channel with a conductance (in symmetrical 140mm KCl solutions) of 210 pS that was activated by noncyclic purine nucleotides and closed by H⁺ ions (pH=6.8); a 211 pS channel that was Ca²⁺-activated and voltage dependent; a 185 pS channel that was blocked by TEA but was insensitive to quinine or nucleotides; a 130 pS channel that was activated by membrane hyperpolarization; and a small conductance (15 pS) channel that was not obviously affected by any manipulation. As determined by radioimmunoassay, cells from passage number 56 secreted 917±128 ng/mg cell protein/48 hr of insulin. In contrast, cells from passage number 77 revealed either no channel activity or an occasional nonselective channel, and secreted only 29.4±8.5 ng/mg cell protein/48 hr of insulin. The nonselective channel found in the passage 77 cells had a conductance of 25 pS in symmetrical 140mm KCl solutions. Thus, there appears to be a correlation between the presence of functional K⁺ channels and insulin secretion.     

Backwater areas as nursery habitats for fishes in pool 13 of the Upper Mississippi River

Samples of larval and juvenile fishes were collected at two depths weekly during spring and summer 1983 near the mouths of backwater areas in Pool 13 of the Upper Mississippi River. The study was conducted to determine the relative value of these habitats as nursery areas for fishes present and to note any interactions that might occur between the backwaters which are being rapidly lost to siltation, and the main channel. The larvae and juveniles collected represented 13 families divided into 27 lower taxa. Cyprinidae, Clupeidae, and Sciaenidae made up 90% of the total catch. Both larvae and juveniles were more abundant near the surface than near the bottom. Densities differed greatly among the three backwater areas studied. Larval fishes were grouped on the basis of their relative abundance in the backwaters or main channel. Overall, more larvae were captured in the backwaters than in main-channel habitats, indicating that backwaters were more productive. In the main channel, densities were greater downstream from the mouths of the backwaters than upstream-possibly indicating that (1) larval fish drifted out of the backwater areas, (2) water rich in nutrients or zooplankton that flowed into the main channel created productive downstream sites that were used as nursery areas, or (3) adult fishes selected downstream sites as spawning areas. Juvenile forms were more abundant in the backwater areas then in the main-channel habitats, some bottom-dwelling fishes excepted. The backwater areas were judged to be important nursery areas for larval and juvenile fishes, and seemed to benefit downstream main-channel sites. Any loss of these habitats would be detrimental to the Mississippi River as a whole.     

Contributions to the mathematical biophysics of branched circulatory systems

A derivation is given of the reflection coefficient of pressure waves in a vessel whose end branches into many smaller vessles. This coefficient depends on the number of these smaller vessels and their sizes relative to the size of the main vessel. Estimations are made of the order of magnitude of the coefficient. Assuming the main vessel to be of the order of size of an artery, it is shown that the reflection coefficient has a value close to one for reflections at branchings into vessels of arteriolar size. It is pointed out that the result may support the idea that the standing waves in the arterial system are due to reflections at the site of the arterioles.     

Sedimentation modified by wind induced resuspension in a shallow tropical lagoon (Cote d'Ivoire)

In shallow environments, under certain conditions of fetch, wind velocity, bathymetry and bottom characteristics, resuspension can be generated by wind induced waves. In the tropical Ebrié lagoon, austral trade winds are dominant almost all year long, and their velocity shows a marked diel pattern with maximum speed between noon and midnight. Only austral trade winds with a speed >3 m s⁻¹ allow particle resuspension which is effective for depths<1.5 m. In these areas, significantly higher values of chlorophyll biomass and mineral seston are noted during the windy sequences. Granulometric and mineralogical analyses showed that only the surficial sediment (0–3 cm) was involved in resuspension. This process induces several effects: 1) an increase of the suspended matter concentration in the water and thus a light attenuation due to a higher turbidity, 2) a redistribution in the whole water column of nutrients from the pore water and 3) a removal of the finer fractions from the superficial sediment. On the contrary, for depths>1.5 m, particle sinking is permanent in depressions which are spontaneously transformed into anoxic systems. At the lagoon scale, sedimentation is significantly modified by wind induced resuspension. According to the bathymetry and the distance from a river, three sedimentary facies are recognized. Their grain size distributions are parabolic in areas where resuspension occurs, logarithmic in areas where no resuspension is possible and hyperbolic in the hollows and the main channels. Finally, a large part of the allochthonous inputs (from drainage and rivers) and autochthonous pelagic production is trapped into the Ebrié lagoon and less than 10% of the particles entering the lagoon are exported toward the Atlantic Ocean.     

A new apparatus for the direct measurement of the effects of otter trawling on benthic nutrient releases

An innovative Towed Trawl Simulator Sledge (TTSS) which has been tested in the field is described. The new instrument, which incorporates six water sampling bottles and video observation equipment, is designed to simulate the effects of the passage of otter trawl groundropes over seafloor sediments. The TTSS is towed from a surface vessel at speeds similar to those of commercial trawls and the water bottles, triggered simultaneously by electrical or acoustic signals, collect water samples in the plume of disturbed sediment behind the groundrope. The TTSS has been tested in the continental shelf fishing grounds of Heraklion Bay (Eastern Mediterranean) where quantitative measurements of the physical (sediment resuspension) and chemical (nutrient release) effects on surface sediments caused by the passage of otter trawl groundrope typical of that used in the Cretan fishery trawling were performed. The performance of the TTSS is also validated against an instrumented full-scale trawl. It is suggested that the TTSS should be used for the study and modelling of the effects of otter trawling and similar fishing processes on nutrient supply from seafloor sediments and the resulting ecosystem responses.     

Comparative use of side and main channels by small‐bodied fish in a large,unimpounded river

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Relative abundance of macroinvertebrates found in habitats associated with backwater area confluences in Pool 13 of the Upper Mississippi River

Ephemeroptera showed the greatest diversity and Diptera (due mainly to Chaoboridae) the greatest absolute abundance among the macroinvertebrate taxa collected in and near three backwater areas of the Upper Mississippi River from 24 April to 24 August 1983. Of the 32 taxa identified, all were insect larvae, pupae, or adults except for one amphipod and one copepod. Three peaks in total abundance were observed — a major one in late July and August and smaller ones in May and June. These peaks corresponded to periods of high temperatures and low flows. When the macroinvertebrates were divided into groups, depending on whether they were found predominantly in the backwater areas, in the main channel, or in a combination of the two, total densities were greater in the backwater areas than in either of the main-channel habitats. In the two main-channel habitats, densities were slightly greater downstream than upstream from the confluences of the backwaters with the main channel, possibly indicating that macroinvertebrates were drifting out of the backwater areas or that nutrients and zooplankton were drifting out of the backwater areas, increasing productivity in downstream areas.     

Anatomy of herpes simplex virus DNA. III. Characterization of defective DNA molecules and biological properties of virus populations containing them.

We have characterized the virus progeny and its DNA from plaque-purified and undiluted passages of herpes simplex virus 1 in HEp-2 cells. Secifically, (i) infectious virus yields declined progressively in passages 1 through 10 and gradually increased at passages 11 through 14. The yields correlated with PFU/particle ratios. (ii) In cells infected with virus from passages 6 through 10, there was an overproduction of an early viral polypeptide (no. 4) and a delay in the synthesis of late viral proteins. In addition, the virus in these passages interfered with the replication of a nondefective marker virus. Cells infected with passage 14 virus produced normal amounts of polypeptide 4 and, moreover, this virus showed minimal interfering capacity. (iii) In addition to DNA of density 1.726 g/cm-3, which was the sole component present in viral progeny of passage 0, passages 6 through 14 contained one additional species (p 1.732) and in some instances (passages 6 and 10) also DNA of an intermediate buoyant density. The ratio of p 1.732 to p 1.726 DNA increased to a maximum of 4 in passages 6 through 9 and gradually decreased to 1 in passages 10 through 14. (iv) p 1.732 DNA cannot be differentiated from p 1.726 DNA with respect to size; however, it has no Hin III restriction enzyme cleavage sites and yields only predominantly two kinds of fragments with molecular weights of 5.1 x 10-6 and 5.4 x 10-6 upon digestion with EcoRI enzyme. (v) Partial denaturation profiles of purified p 1.732 DNA from passage 14 revealed the presence of two types of tandemly repeated units corresponding roughly in size to the EcoRI fragments and situated in different molecules. (vi) In addition to the two kinds of p 1.732 molecules consisting of tandem repaeat units of different sizes, other evidence for the diversity of defective DNA molecules emerged from comparisons of specific infectivity and interfering capacity of the progeny from various passages. The data suggest that some of the particles with DNA of normal buoyant density (1.726) must also be defective since the capacity to interfere and to produce an excess of polypeptide 4 did not appear to be proportional to the amount of high-buoyant-density defective DNA. The data suggest that defective interfering particles are replaced by defective particles with diminished capacity to interfere and that more than one species of defective DNA molecules evolves on serial preparation of HSV.     

Measurement of resuspended aerosol in the Chernobyl area

Results of measurements of the resuspended radioactive aerosols in the Chernobyl area are presented which were obtained soon after the Chernobyl reactor accident and in a European project in 1992–1993. The measurements were carried out with the intention of obtaining a data base for dose assessment of resuspended radioactive particles. Potential significant dose contributions may result from inhalation and secondary contamination due to resuspended radionuclides. In this first article of a series of three papers, the instrumentation and the measurement uncertainties are discussed. An effort was made to sample quantitatively giant aerosol particles (particles larger than 10 μm aerodynamic diameter) as well. The comparison of the samplers shows, in general, an agreement of concentration measurements of ¹³⁷Cs and ⁷Be within a factor of two. One sampler was identified with larger discrepancies and needs additional investigation of its sampling characteristics; for another device, the recalibration of the analysing system is recommended. Ordinary integrating samplers have a loss of about 30% in ¹³⁷Cs activity compared to an isokinetic sampler collecting giant particles as well. The mean ratio of ¹³⁷Cs activity concentration between an instrument sampling only particles larger than 10 μm and an ordinary integrating sampler is 0.39 ± 0.15 during anthropogenic-enhanced resuspension. These findings demonstrate the significant contribution of giant particles to resuspended airborne radioactivity. The results of this study concerning integral measurements during wind-driven resuspension proved to be in good agreement with previously published data on resuspension. Received: 15 May 1997 / Accepted: 4 August 1997