The influence of phytoplankton composition on the relative effectiveness of grinding and sonification for chlorophyll extraction

The chlorophyll recovery efficiency was compared between control, ground, and sonified samples. The results showed significant improvement between control and ground samples but not between control and sonified samples. Neither prolonging time of sonification nor using an ice bath during filter grinding improved efficiency. Higher chlorophyll a recovery was obtained from ground samples than from sonified ones, when the water samples contained centric diatoms and filamentous blue-green algae. When total phytoplankton numbers were high, there was a distinct advantage in using grinding rather than sonification for chlorophyll c recovery.Contribution no. 313 of the Great Lakes Research Division, University of Michigan.Contribution no. 313 of the Great Lakes Research Division, University of Michigan.     

A comparative ultrastructural and morphometric study of six species of the diatom genus Stephanodiscus

The cellular composition of six species of Stephanodiscus commonin the Laurentian Great Lakes was compared using quantitativeultrastructural and morphometric techniques. The species studiedspan a considerable portion of the range of size and obviousmorphological differences known within the genus. Results showconsiderable differences between species in the amount of totalcell volume occupied by frustule, cytoplasm, vacuole and chloroplast.Other organelles, such as mitochondria, Golgi bodies and thenucleus occupy similar fractions of the total cytoplasmic massin all species studied. The main within-species difference isfound in the amount of storage products present, which appearsto be related to cell condition. The techniques utilized areapplicable to populations in mixed assemblages from the fieldand can be used to measure the cellular composition of suchpopulations with a high degree of accuracy and precision. ^*Contribution No. 406 of the Great Lakes Research Division,The University of Michigan.     

Primary productivity and nutrients in the sediment retention basin of Lake Monroe

Lake Monroe is the largest body of water in Indiana with a daily mean productivity of 220 mg · C · m⁻² · day⁻¹ in an observed range from 26 to 714 m · C · M⁻² day⁻¹. It is a medium soft reservoir; the acid combining capacity varies from 0.28 to 0.71 meq · l⁻¹ with a mean slightly above 0.5 meq · l⁻¹. The results of diurnal changes in major nutrients, the C, N, and P ratios, and bioassay experiments indicated that phosphorus is the major limiting nutrient on algal photosynthesis in this lake. Surface photo-inhibition may be used to indicate the sufficiency of light for the species of algae in the water. The low productivity in December, January, February, and early March can be attributed to light limitation due to low water transparency. Contribution no. 312 of the Great Lakes Research Division, University of Michigan. Contribution no. 312 of the Great Lakes Research Division, University of Michigan.     

In situ predatory behavior ofMysis relicta in Lake Michigan

Selectivity coefficients (W) and predation rates on Lake Michigan zooplankton were determined forMysis relicta during spring through fall using anin situ method. W values indicated the following ranked order of prey preference: Cladocera > copepod copepodites and copepod nauplii > adult diaptomids and cyclopoids. With few exceptions, W values for different prey categories remained fairly constant despite greatly changing relative abundances of prey. Predation rates and prey selectivity were similar in most cases to those determined in laboratory studies. Ingestion rates (percent dry body weight · day^–1) were correlated to total prey biomass (r = 0.38) and to effective prey biomass (r = 0.85), where the weighting factors were overall mean selectivity coefficients for the different prey categories. This result suggested that seasonally varying composition of prey caused much of the variation in ingestion rates among experiments. Feeding trials performed at the same depth with daytime and nighttime assemblages of zooplankton indicated that Cladocera may escape heavyMysis predation at night by migrating from the metalimnetic-hypolimnetic interface into the epilimnion.Contribution 333 from the Great Lakes Research Division, The University of Michigan and contribution 287 from the Great Lakes Environmental Research Laboratory, National Oceanic and Atmospheric Administration.Contribution 333 from the Great Lakes Research Division, The University of Michigan and contribution 287 from the Great Lakes Environmental Research Laboratory, National Oceanic and Atmospheric Administration.     

A numerical simulation of trichromatic equations in chlorophyll estimation using the spectrophotometric technique

A numerical simulation of trichromatic pigment equations is made with the aid of a computer utility program. Significant quantitative differences in the estimates of pigment concentration result from using different sets of trichromatic equations. Estimates of chlorophylls a, b, and c were found highly correlated with the application of the equations, even though the absorbance values used as input for the stimulation are not correlated.Contribution No. 284 of the Great Lakes Research DivisionContribution No. 284 of the Great Lakes Research Division     

Phytoplankton dynamics within the subsurface chlorophyll maximum of Lake Michigan

The Lake Michigan deep chlorophyll maximum during July 1977was a broad band of phytoplankton –15 m thick rather thanthe thin layers observed in smaller lakes. The maximum concentrationof chlorophyll was typically four times surface values. ¹⁴Cuptake incubations showed that the deep plankton layer was aviable component of the pelagic ecosystem accounting for upto 60% of primary production on an areal (m²) basis. The LakeMichigan chlorophyll maximum was composed primarily of largediatoms, filaxnentous green algae, and flagellates. Althoughlarge scale physical processes such as seiches had some effecton the chlorophyll maximum, the basic integrity of the layerwas maintained by biological processes. In particular, epilinmeticnutrient depletion was apparently a major factor affecting thevertical distribution of phytoplankton. Application of a diffusionmodel to the chemical and chlorophyll data indicated that nutrientuptake and primary production by phytoplankton were more importantin determining chemical and chlorophyll vertical profiles thandiffusivity. ¹Contribution No. 389 from the Great Lakes Research Division,University of Michigan.     

The release of dissolved organic carbon as a result of diatom fragmentation during feeding byMysis relicta

The release of dissolved organic carbon (DOC) byMysis relicta that occurs while feeding may be an important pathway in the recycling of nutrients in Lake Michigan. Laboratory experiments were conducted to determine the levels of DOC released byM. relicta from Lake Michigan while grazing on laboratory-cultured diatoms. Increasing filtering rates were observed at progressively lower chlorophyll concentrations. Higher filtering rates were accompanied by higher fragmentation of diatoms and a higher concomitant release of DOC from broken cells. While the amount of DOC release is small (0.02–3.30 g C · 1^–1· h^–1), it nonetheless is of importance to bacteria and possibly some algae. After assimilation by bacteria and algae, the organic carbon is then available to consumers.Contribution No. 239, Center for Great Lakes Studies, The University of Wisconsin-Milwaukee, Milwaukee, WI 53201, U.S.A.Contribution No. 239, Center for Great Lakes Studies, The University of Wisconsin-Milwaukee, Milwaukee, WI 53201, U.S.A.     

Xenopus laevis larvae (Amphibia,Anura) as model suspension feeders

Larvae of the South African clawed frog, Xenopus laevis (Daudin), are efficient, obligate suspension feeders. We examine the relationship between the ambient particle concentration offered these larvae as food and their filtering, ingestion, and buccal pumping rates. We demonstrate that: (i) the larvae can sense and respond to a broad range of particle concentrations, down to 0.2 mg 1^–1 (dry weight); (ii) their metabolic needs theoretically can be met by particle concentrations as low as 5 mg 1^–1; and (iii) their patterns of regulation of filtering and ingestion fit predictions from certain models used to describe zooplankton feeding dynamics. Two such models are discussed: the modified Monod (Michaelis-Menten) model, with a lower threshold below which the tadpoles do not feed, and an energy optimization model. Both the models and the observed behavior of the tadpoles allow for stability of populations of food organisms. Tadpole feeding dynamics apparently are compatible with both the predictions and assumptions of these models, suggesting similar regulation of feeding by tadpoles and zooplankton. However, the size, morphology, and behavior of X. laevis larvae make their feeding regulation uniquely accessible to direct observation.Contribution No. 223, Center for Great Lakes Studies, University of Wisconsin-Milwaukee, Milwaukee, Wisconsin.Contribution No. 223, Center for Great Lakes Studies, University of Wisconsin-Milwaukee, Milwaukee, Wisconsin.     

Population genetic structure of the round goby in Lake Michigan: implications for dispersal of invasive species

Understanding subsequent dispersal of non-native species following introduction is important for predicting the extent and speed of range expansion and is critical for effective management and risk assessment. Post-introduction dispersal may occur naturally or via human transport, but assessing the relative contribution of each is difficult for many organisms. Here, we use data from seven microsatellite markers to study patterns of dispersal and gene flow among 12 pierhead populations of the round goby (Neogobius melanostomus) in Lake Michigan. We find significant population structure among sampling sites within this single Great Lake: (1) numerous populations exhibited significant pairwise F _ST and (2) a Bayesian assignment analysis revealed three distinct genetic clusters, corresponding to different pierhead locations, and genetic admixture between these clusters in the remaining populations. Genetic differentiation (F _ST) is generally related to geographic distance (i.e., isolation by distance), but is periodically interrupted at the scale of Lake Michigan due to gene flow among geographically distant sites. Moreover, average genetic differentiation among populations exhibit a significant, negative correlation with the amount of shipping cargo at ports. Our results, therefore, provide evidence that genetic structure of the round goby in Lake Michigan results from limited natural dispersal with frequent long-distance dispersal through anthropogenic activities such as commercial shipping. Our study suggests that while round gobies can undoubtedly disperse and found new populations through natural dispersal mechanisms, their spread within and among the Great Lakes is likely aided by transport via ships. We, therefore, recommend that ballast-water treatment and management may limit the spread of non-native species within the Great Lakes after the initial introduction in addition to preventing the introduction of non-native species to the Great Lakes.     

Modelling local and long-distance dispersal of invasive emerald ash borer Agrilus planipennis (Coleoptera) in North America

Limiting the damage by non-indigenous species requires rapid determination of current and potential distributions and vectors of dispersal, and development of appropriate management measures. The emerald ash borer ( Agrilus planipennis ), a wood-boring beetle native to South-East Asia, was first reported in the Great Lakes region during summer 2002. The beetle poses an enormous threat to native ash ( Fraxinus ) species of North America, as untreated trees in infested areas of Ontario, Michigan and Ohio suffer high mortality. We demonstrate that the borer has spread in North America through a combination of diffusive range extension, associated with local flights, and by long-distance 'jump' dispersal associated with human movement of infested sapling or contaminated firewood. Probability of infestation was inversely related to distance from borer epicentres but positively related to the size of human population centres. At least 9 of 39 populations that were first reported in Michigan during 2004 cannot be accounted for by local diffusion, raising the possibility that other unidentified mechanisms may be contributing to the dispersal of the beetle. In the absence of quarantine, by 2005 all of Michigan's lower peninsula was contained within the boundaries of potential diffusive range expansion. Infested ash saplings also were introduced from Michigan to Maryland during 2003, and subsequently transplanted to five sites in Maryland and Virginia. Quarantine and eradication measures have had mixed results: in the south-central USA, the species appears on the brink of eradication, whereas its distribution has continued to spread during 2005 in the Great Lakes region despite extensive containment and quarantine measures. Quarantine success in the Great Lakes region is encumbered by multiple dispersal vectors, larger borer population sizes and by the more extensive geographical distribution that was achieved prior to implementation of control measures.     

Stratigraphic and dynamic effects of sediment reworking by Great Lakes zoobenthos

Recent sediments of the North American Great Lakes are inhabited by numerous species of macrobenthos which alter the physical and chemical properties of sediments and modify interface transport characteristics. Distributions of such radionuclides as cesium-137, lead-210, and isotopes of plutonium exhibit a zone of constant activity extending down from the sediment-water interface from 1 to 15 cm. Recent studies have confirmed that radiometrically determined mixed depths are consistent with the vertical distribution of oligochaete worms and the amphipod,Pontoporeia hoyi. Generally, 90% of the benthos are contained within the radiometrically defined mixed zone. Where comparisons are possible, rates of sediment reworking by ‘conveyor belt’ species are comparable to or exceed sedimentation rates. Systematic variations in the mixed depth occur within depositional basins with greatest depths tending to be associated with least consolidated, organically rich materials. A quantitative steady-state mixing model accounts satisfactorily for observed radioactivity and heavy metal profiles. Bioturbation appears to be an important process, limiting the resolution with which historical records of particle-associated contaminants may be reconstructed from sediment cores. As bioturbation serves to maintain contact of contaminated sediments with overlying water, this time may also characterize the long-term lake recovery for contaminants removed by burial. As the time varies with location, a mean for an entire lake is not well known, but is on the order of 20 years for Lake Huron. Contribution No. 300 of the Great Lakes Environmental Research Laboratory, Ann Arbor, Michigan.     

Patterns and Pathways in the Post-Establishment Spread of Non-Indigenous Aquatic Species: The Slowing Invasion of North American Inland Lakes by the Zebra Mussel

The zebra mussel, Dreissena polymorpha, has spread through eastern North American aquatic ecosystems during the past 15 years. Whereas spread among navigable waterways was rapid, the invasion of isolated watersheds has progressed more slowly and less predictably. We examined the patterns of overland spread over multiple spatial and temporal extents including individual lake districts, states, and multi-state regions in the USA and found that only a small proportion (<8%) of suitable inland lakes have been invaded, with the rate of invasion appearing to be slowing. Of the 293 lakes known to be invaded, 97% are located in states adjacent to the Laurentian Great Lakes with over half located in Michigan. Only six states have more than 10 invaded lakes and only in Michigan and Indiana have more than 10% of suitable lakes become invaded. At smaller spatial extents, invaded lakes are often clustered within a lake-rich region across southern Michigan and northern Indiana. This clustering appears primarily due to multiple overland invasions originating from the Great Lakes followed to a lesser extent by subsequent secondary overland and downstream dispersal. Downstream spread appears responsible for only one third of the inland invasions. Temporally, invasions peaked in the late 1990s, with only 13 new invasions (0.4% of suitable lakes) reported in 2003 in the four-state region surrounding Lake Michigan. Peak rates of invasion occurred 4–6 years earlier in Michigan relative to Indiana and Wisconsin, but this time lag is likely due to differences in the establishment of Great Lake source populations rather than ‘stepping stone’ dispersal across the landscape.     

Clostridium botulinum type E in fish from the Great Lakes

Bott, Thomas L. (University of Wisconsin, Madison), Janet S. Deffner, Elizabeth McCoy, and E. M. Foster. Clostridium botulinum type E in fish from the Great Lakes. J. Bacteriol. 91:919-924. 1966.-The intestinal contents of more than 3,000 fish from Lakes Erie, Superior, Huron, and Michigan were examined for Clostridium botulinum type E. Demonstration of the organism was accomplished by identifying its toxin in liquid cultures inoculated with material from the alimentary tract. Incidence figures, expressed as per cent of the fish tested, were: Lake Erie, 1%; Lake Superior, 1%; Lake Huron, 4%; the main body of Lake Michigan, 9%; and Green Bay (on Lake Michigan), 57%. Thus, C. botulinum type E appears to be widely but unevenly distributed in the Great Lakes, and fish from all areas are potential carriers of it.     

Influence of experimental conditions on nitrogenous excretion by Lake MichiganMysis relicta (Lovén): laboratory studies with animals acclimated inFragilaria

In laboratory experiments, rates of excretion of ammonia and urea by Lake MichiganMysis relicta were compared for animals incubated in the presence and the absence of algal food (Fragilaria crotonensis chemostat outflow). Prior to experiments, all animals were acclimated to laboratory conditions and the experimental food for 2–4 weeks. Algae used in experiments were enriched in the dark with nutrients (N and P) prior to experimental incubation.There were no significant differences in the ammonia and urea excretion rates of mysids incubated in filtered water compared to those inFragilaria cultures, or in ammonia excretion rates either as a function of individual size or sex of the mysid or of its having been held at 5 °C, 10 °C, or 15 °C. Ammonia excretion rates measured between 4 and 8 h of incubation were significantly higher than those between 1–3 h and 8–15 h, for mysids held both in filtered water and inFragilaria cultures. The results are compared to those from shipboard incubations in a previous study and are discussed with respect to physiological regulation of nitrogenous waste production.Contribution No. 234, Center for Great Lakes Studies, University of Wisconsin-Milwaukee, Milwaukee, Wisconsin.Contribution No. 234, Center for Great Lakes Studies, University of Wisconsin-Milwaukee, Milwaukee, Wisconsin.     

Response of bacteria and phytoplankton to contaminated sediments from Trenton Channel,Detroit River

Several types of bioassays were used in 1986 and 1987 to investigate the effect of contaminated sediments on natural populations of bacteria and phytoplankton from the Trenton Channel, Detroit River. The approach included the measurement of uptake of ³H-glucose or ³H-adenine by bacteria and ¹⁴C-bicarbonate by phytoplankton in the presence of different amounts of Trenton Channel and Lake Michigan (control) sediments. Trenton Channel sediments are contaminated by high levels of toxic organic compounds and metals, especially zinc, lead, and copper. Because levels of biomass of bacteria and phytoplankton varied widely among the different bioassays, it was necessary to adjust uptake rates for biomass. Biomass adjustments were made using acridine orange counts for bacteria and chlorophyll measurements for phytoplankton. The results show a statistically significant suppression of uptake of substrates for both bacteria and phytoplankton with increasing amounts of sediment. Uptake was suppressed as much as 90 percent for bacteria and 93 percent for phytoplankton at 1200 mg l^-1 of Trenton Channel sediments compared to bioassays without sediment. Uncontaminated Lake Michigan sediment suppressed uptake much less than Detroit River sediment; the difference in suppression of uptake between the two types of sediment was statistically significant for both bacteria and phytoplankton.Contribution No. 518 of the Center for Great Lakes and Aquatic Sciences of the University of Michigan.     

Density-dependent dispersal in birds and mammals

Density‐dependent dispersal can be caused by various mechanisms, from competition inducing individuals to emigrate (positive density‐dependence) to social crowding effects impeding free movement (negative density‐dependence). Various spatial population models have incorporated positively density‐dependent dispersal algorithms, and recent theoretical models have explored the conditions for density‐dependent dispersal (DD) to evolve. However, while the existence of DD is well documented in some taxa such as insects, there is no clear picture on its generality in vertebrates. Here I review the available empirical data on DD in birds and mammals, focusing mainly on variation in dispersal between years and on experimental density manipulations. Surprisingly few studies have explicitly focused on DD, and interpretation of the available data is often hampered by differences in approach, small sample sizes and/or statistical shortcomings. Positive DD was reported in 50 and 33% of the selected mammal and bird studies, respectively, while two studies on mammals (out of eight) reported negative DD. Among bird studies, DD was more often reported for emigration rates or long‐distance recoveries than for average distances within finite study areas. Experimental studies manipulating densities (mainly on mammals) have consistently generated positive DD, typically showing reduced emigration in response to partial population removal. Studies that examined dispersal in relation to seasonal changes in density (small mammals only) have more often reported negative DD. Studies that compared dispersal between sites differing in density, also show a mixture of positive and negative DD. This suggests that dispersal changes in a more complex way with seasonal and spatial density variation than with annual densities, and/or that these results are confounded by other factors differing between seasons and sites, such as habitat quality. I conclude that both correlational and experimental studies support the existence of positive, rather than negative, density‐dependent dispersal in birds and mammals.     

A centrifugation technique for separating ascospores from soil

Summary A centrifugation technique employing sucrose density gradient columns has been developed for separating fungus spores from soil mineral particles. Four pyrenomycetes plus a number of sterile isolates have been obtained by the use of this technique. The technique may be applicable to other types of fungi producing large, readily identifiable spores.Contribution from the Soil and Water Conservation Research Division, Agricultural Research Service, USDA, in cooperation with the Washington Agricultural Experiment Station. Scientific Paper 2830, Washington State University, College of Agriculture.     

The physics of sediment transport,resuspension, and deposition

The physics of sediment transport, resuspension and deposition is reviewed. First, the general problem is described and then emphasis is given to recent quantitative research in Lakes Ontario, Michigan and Erie. In this discussion, some original calculations are presented to show that Ekman layer sediment transport is important in determining the deposition areas in deep lakes.GLERL Contribution No. 436     

Use of ecological indicators to assess the quality of Great Lakes coastal wetlands

Over 2000 coastal wetland complexes have been identified in the Laurentian Great Lakes watershed, each providing critical habitat for numerous aquatic and terrestrial species. Research has shown there is a direct link between anthropogenic activities (urbanization and agricultural development) and deterioration in wetland health in terms of water quality and biotic integrity. In this study, we evaluate coastal marshes throughout the Great Lakes basin using a suite of published ecological indices developed specifically for coastal wetlands of the Great Lakes (Water Quality Index (WQI), Wetland Macrophyte Index (WMI), and the Wetland Fish Index (WFI_Basin)). We surveyed 181 wetlands, including 19 in Lake Superior (11%), 11 in Lake Michigan (6%), 13 in Lake Huron (7%), 92 in Georgian Bay and the North Channel (51%), 18 in Lake Erie (10%), and 28 in Lake Ontario (15%), over a 13 year period (1995–2008). Water quality parameters were measured at every site, while paired fyke nets were used to assess the fish community (132 sites) and macrophytes were surveyed and identified to species (174 sites); all of this information was used to calculate the associated index scores. One-way ANOVA results showed that there were significant differences in wetland quality among lakes. According to the WQI, we found that over 50% of marshes in Lakes Michigan, Erie, and Ontario were in degraded condition, while over 70% of marshes in Lakes Superior, Huron, and Georgian Bay were minimally impacted. Georgian Bay had the highest proportion of wetlands in very good and excellent condition and least number of wetlands in a degraded state. The WMI and WFI showed similar results. This is the largest bi-national database of coastal wetlands and the first study to provide a snapshot of the quality of coastal habitats within the Great Lakes basin. We recommend this information be used to guide conservation and restoration efforts within the Laurentian Great Lakes.     

Predaceous feeding habits of Limnocalanus macrurus

Limnocalanus macrurus, a large, glacial-relict copepod, hasbeen assumed an omnivore or a herbivore; predaceous habits ofthe species are unknown. The predaceous feeding habits of Limnocalanusfrom Lake Michigan were studied in the laboratory using naturalprey. Predation rates were highest on copepod nauplii. Copepoditesof Diaptomus spp. and Cyclops spp. were preyed upon at lowerrates. Limnocalanus preyed selectively upon nauplii <300µm. Small cyclopoid copepodites (<–750 µm)were also selected over large copepodites. Experiments usingtwo prey types showed that nauplii were selected over all copepodites,and that no selectivity existed for either diaptomid or cyclopoidcopepodites. Predaceous feeding habits began in the fourth copepoditestage of Limnocalanus. Predaceous feeding rates of Limnocalanuschanged seasonally being highest in late spring and autumn andlowest in summer and early winter. Since Limnocalanus also feedson net-phytoplankton, predation rate changes may be relatedto changes in the relative abundance of large phytoplanktonand naupliar prey in nature. Limnocalanus predation may be animportant factor in structuring the zooplankton community. Present address: Great Lakes Research Division, University Michigan,Ann Arbor, MI 48109, USA