Growth rate and longevity of Dreissena polymorpha (Pallas): a review and recommendations for future study.ABSTRACT We review the variety of methods that have been used over the last 50 y in the Former Soviet Union, Eastern and Western Europe Western Europe
The countries of western Europe, especially those that are allied with the United States and Canada in the North Atlantic Treaty Organization (established 1949 and usually known as NATO). , and recently in North America North America, third largest continent (1990 est. pop. 365,000,000), c.9,400,000 sq mi (24,346,000 sq km), the northern of the two continents of the Western Hemisphere. to determine growth rate and longevity in zebra mussels zebra mussel
Either of two species of tiny mussels (genus Dreissena) that are prominent freshwater pests. They proliferate quickly and adhere in great numbers to virtually any surface. (Dreissena polymorpha Noun 1. Dreissena polymorpha - inch long mollusk imported accidentally from Europe; clogs utility inlet pipes and feeds on edible freshwater mussels
zebra mussel [Pallas]). These methods include: counting annual rings annual rings, the growth layers of wood that are produced each year in the stems and roots of trees and shrubs. In climates with well-marked alternations of seasons (either cold and warm or wet and dry), the wood cells produced when water is easily available and , analysis of size-frequency distributions, following growth under experimental conditions and monitoring marked mussels under natural conditions, without removing them from substrate. The last method provides the most reliable data, however this is the least common method used. Dreissena polymorpha growth rates Growth Rates
The compounded annualized rate of growth of a company's revenues, earnings, dividends, or other figures.
Remember, historically high growth rates don't always mean a high rate of growth looking into the future. depend on water temperature, season of the year, location in the water column, food availability, oxygen concentrations, water velocity and various other environmental factors. However, it is very difficult to separate the independent effects of each of these factors, especially in natural waterbodies. Several factors may overlap and have additive or synergistic effect Synergistic effect
A violation of value-additivity in that the value of a combination is greater than the sum of the individual values. that makes it difficult to determine the effects of a single factor. When comparing among studies that used the same methods, we found that zebra mussels grow faster in reservoirs than in lakes. The reported longevity of D. polymorpha varies from 2-19 y and it is not clear to what extent this variation is caused by biological variability biological variability Lab medicine The variability in a lab parameter due to physiologic differences among subjects–interindividual BV, and in the same subject over time–intraindividual BV and environmental conditions and what amount of the variation is caused by the methods used to assess age and longevity.
KEY WORDS: zebra mussels, Dreissena polymorpha, growth, growth rate, methods, longevity
The zebra mussel, Dreissena polymorpha (Pallas), is one of the most pervasive invaders in freshwaters of the northern hemisphere. However, many aspects of the basic biology of D. polymorpha that are necessary for understanding and predicting the population dynamics Population dynamics is the study of marginal and long-term changes in the numbers, individual weights and age composition of individuals in one or several populations, and biological and environmental processes influencing those changes. and ecological impacts of this invader are still not well known. In addition, much of the research on the biology of D. polymorpha that has been conducted in the former Soviet Union (FSU FSU Florida State University
FSU Former Soviet Union
FSU Ferris State University
FSU Fayetteville State University (North Carolina)
FSU Frostburg State University
FSU Finance Sector Union ), has not been published in English, and therefore it is not available to most scientists currently studying D. polymorpha. Growth rate and longevity are particularly important for understanding the population biology Population biology is a study of biological populations of organisms, especially in terms of biodiversity, evolution, and environmental biology.
Malthus can almost be considered an early population biologist, even though his training was in economics and the term population and ecological impacts of zebra mussels, especially because fecundity fecundity /fe·cun·di·ty/ (fe-kun´dit-e)
1. in demography, the physiological ability to reproduce, as opposed to fertility.
2. ability to produce offspring rapidly and in large numbers. and filtering capacity increase with body size. Most of the published research has been conducted with the invasive subspecies subspecies, also called race, a genetically distinct geographical subunit of a species. See also classification. Dreissena polymorpha polymorpha, which is capable of living in totally fresh water and has been the major invader in most places where dreissenids have been introduced. Less work has been conducted with Dreissena bugensis (Zhuravel 1951, MacIsaac 1994, Baldwin et al. 2002), Dreissena polymorpha andrusovi (Karpevich 1952, 1964, Lvova et al. 1983, 1994) and Dreissena caspia (Karpevich 1952, 1964).
Here we review the variety of methods used to estimate growth and longevity of zebra mussels over the last 50 y (Table 1, Table 2), discuss limitations of each and recommend the most appropriate methods for measuring growth and longevity in the field. We also synthesize To create a whole or complete unit from parts or components. See synthesis. the impacts of a range of environmental factors on growth and longevity in zebra mussels.
METHODS TO ESTIMATE GROWTH RATE
Rings on Shells
One of the oldest and most common methods for estimating the growth rate of zebra mussels is by counting annual rings on shells of different sizes, and then calculating the average length of each age group of Dreissena in a population (Karpevich 1952, 1964, Kachanova 1963, Stanczykowska 1964, Lyakhov & Mikheev 1964, Mikheev 1964, Kornobis 1977, Karatayev & Tishchikov 1979, Kirpichenko & Antonov 1982, Dorgelo & Gorter 1984, Draulans & Wouters 1988, Miroshnichenko 1990). Plotting the average size of each age group against their age provides a growth rate curve. The advantage of this method is that by measuring individuals at a single point in time estimates of growth over several years can be made. However, counting growth rings is very subjective as it is difficult to distinguish annual rings from rings formed because of other factors that slow growth. Morton (1969a) found that two rings are formed annually: when growth slows during the winter and during spawning. Lvova (1980) found in the Uchinskoe Reservoir, 3-9 rings on the shells of 3-y-old mussels grown in cages for 2 y. In Czos Lake Lewandowski (1983) found from 1-3 rings on the shells of 1-y-old D. polymorpha, and from 2-5 rings on 2-y-old mussels.
Many other authors have also reported difficulties in distinguishing annual rings (Karpevich 1964, Kirpichenko 1965, Morton 1969a, Wiktor 1969, Lvova-Kachanova 1972, Lvova 1980, Lewandowski 1982a, Karatayev 1983, bij de Vaate 1991, Lvova et al. 1994). Often mussels with distinct rings can be found side by side with mussels without rings (Lvova 1980, Lvova et al. 1994, Jantz 1996). Moreover, 1-mm zebra mussels that settle at the end of the growing season growing season, period during which plant growth takes place. In temperate climates the growing season is limited by seasonal changes in temperature and is defined as the period between the last killing frost of spring and the first killing frost of autumn, at which do not produce a first annual ring. Therefore, these mussels would he incorrectly identified as young-ofthe-year the following year.
Size-frequency distributions have been used in a number of studies of D. polymorpha growth rates (Morton 1969a, Jantz & Neumann 1992, Martel 1993, 1995, Smit et al. 1993, Dall & Hamburger 1996, Chase & Bailey 1999a, Orlova & Panov 2004), and can be useful if there is highly synchronized syn·chro·nize
v. syn·chro·nized, syn·chro·niz·ing, syn·chro·niz·es
1. To occur at the same time; be simultaneous.
2. To operate in unison.
1. spawning and settlement and low interindividual variation in growth. Newly settled mussels form a distinct size class maintain distinct size structure for all age/size classes (Golikov 1970). However, in many water bodies D. polymorpha spawn To launch another program from the current program. The child program is spawned from the parent program.
(operating system) spawn - To create a child process in a multitasking operating system. E.g. throughout the entire summer, producing several peaks in veliger ve·li·ger
A larval stage of a mollusk characterized by the presence of a velum.
[New Latin v densities during the year (Lvova 1977, 1980, Karatayev 1983, Lvova et al. 1994, Burlakova 1998) and a wide size range (up to 16 mm difference in size) by the end of their first growing season (Wesenberg-Lund 1939, Mikheev 1964, Kirpichenko 1971, Szlauer 1974, Lvova 1977, Lewandowski 1983, Neumann et al. 1993, Martel 1995). Therefore, age classes will not form distinct size classes (Karatayev 1983, bij de Vaate 1991, Jantz & Neumann 1992, Lvova et al. 1994). This method is most effective when D. polymorpha spawn synchronously, have fast growth and are short lived, such as the shallow areas of the Svisloch River (Burlakova 1998). In this river D. polymorpha settle in the summer and grow during the following year, producing two distinct size classes of mussels (0+ and +) because the majority of older mussels die over winter because of fluctuating water levels and predation predation
Form of food getting in which one animal, the predator, eats an animal of another species, the prey, immediately after killing it or, in some cases, while it is still alive. Most predators are generalists; they eat a variety of prey species. by ducks. At deeper depths mussels survive longer, producing many age classes, which are less distinct as cohorts, making this method less useful (Burlakova 1998). This method could also be used for studies that follow growth on experimental substrates when the time of settlement is known (Lvova 1977, Sprung 1992).
Growth Under Experimental Conditions
Many studies have been used to estimate D. polymorpha growth under experimental conditions, especially in cages. In the FSU this method was used in Kuybyshevskoe (Mikheev 1964), Uchinskoe (Lvova-Kachanova 1972, Lvova 1980) and Tsimlyanskoe (Lvova et al. 1983) reservoirs, in Lake Lukomskoe (Karatayev 1983), the Narochanskie lakes and the Svisloch River (Burlakova 1998). More recently, this method has been used by East European (Stanczykowska & Lewandowski 1995), West European (Smit et al. 1992, 1993, Sprung 1992, 1995a, Dorgelo 1993, Dali & Hamburger 1996) and North American North American
named after North America.
North American blastomycosis
see North American blastomycosis.
North American cattle tick
see boophilusannulatus. scientists (Bitterman et al. 1994, MacIsaac 1994, Allen et al. 1999, Horvath & Lamberti 1999, Yu & Culver cul·ver
A dove or pigeon.
[Middle English, from Old English culufre, from Vulgar Latin *columbra, from Latin columbula, diminutive of columba, dove.] 1999, Garton & Johnson 2000).
Other methods for studying mussel mussel, edible freshwater or marine bivalve mollusk. Mussels are able to move slowly by means of the muscular foot. They feed and breathe by filtering water through extensible tubes called siphons; a large mussel filters 10 gal (38 liters) of water per day. growth under experimental conditions include growing mussels on artificial substrates (Dorgelo & Goner gon·er
One that is ruined or doomed.
Slang a person who is about to die or who is beyond help
1984, Sprung 1992, Martel 1993), and in the laboratory (Walz 1978a, 1978b, Jantz & Neumann 1992, 1998, Dorgelo 1993, Neumann et al. 1993, Jantz 1996, Baldwin et al. 2002). All of these methods could produce different types of artifacts artifacts
see specimen artifacts. , which may influence observed growth rates. Mesh, usually 3-5 mm, may prevent normal water flow through the cage, particularly for smaller mesh sizes (e.g., 1.2 mm, Garton & Johnson 2000). Cages can also be overgrown overgrown
said of a part that has not been kept trimmed.
overgrown hooves put unusual stresses on bones and tendons and allow for distortion of the wall and sole. by periphyton pe·riph·y·ton
Sessile organisms, such as algae and small crustaceans, that live attached to surfaces projecting from the bottom of a freshwater aquatic environment. , further reducing water flow (Kachanova 1963, Karatayev 1983, Stanczykowska & Lewandowski 1995, Burlakova 1998). However, the effects of caging artifacts on growth rates are usually not well tested or quantified (see Burlakova 1998 later).
Marked Mussels Under Natural Conditions
Following tagged mussels under natural conditions has been used in very few studies (e.g., Stoeckman & Garton 1997, Burlakova 1998) although this method could provide the most realistic estimates of zebra mussel growth rates. Burlakova (1998) found that the growth rate of mussels on stones in the Svisloch River was greater than that for caged mussels in the same environment (Fig. 1A). Early in the spring (April), when macrophytes and periphyton abundances were low, the difference between caged and uncaged un·caged
1. Not confined in a cage: uncaged birds.
2. Released from a cage: an uncaged lion in the arena. mussels was small (~30%). This difference increased to almost 400% in the middle and especially by the end of the growing season when the quantity of drifting plants in the water increased and periphyton densities were high (Burlakova 1998). The disadvantage of this method is that zebra mussels can move, form druses or be consumed by predators, making it difficult to follow individuals through time.
[FIGURE 1 OMITTED]
IMPACT OF ENVIRONMENTAL FACTORS ON GROWTH RATE
The growth rate of Dreissena depends on water temperature, season of the year, location in the water column, trophic trophic /tro·phic/ (tro´fik) (trof´ik) pertaining to nutrition.
Of, relating to, or characterized by nutrition. conditions, which affect food availability, and water velocity as well as other environmental factors (Table 1).
It is well established that the growth rate of D. polymorpha is accelerated by increased water temperature (Table 1). Especially convincing are data from studies in different temperature zones of cooling water reservoirs for thermal power plants in the FSU (Yaroshenko & Naberezhnyi 1971, Skalskaya 1976a, 1976b, Elagina et al. 1978, Karatayev & Tishchikov 1979, Karatayev 1983, 1984, 1988) and other areas of Eastern Europe Eastern Europe
The countries of eastern Europe, especially those that were allied with the USSR in the Warsaw Pact, which was established in 1955 and dissolved in 1991. (Stanczykowska 1976a, Kornobis 1977). In these studies D. polymorpha growth rates were compared among the various temperature zones of the same waterbody or lakes within same lake system. Therefore, environmental conditions other than temperature were similar, allowing a direct estimate of thermal effects.
However, when the maximum temperature is >30[degrees]C, D. polymorpha growth decreases, and at temperatures >32[degrees]C most mussels die. In the hottest zone of Lukomskoe Lake, where maximum summer temperature exceeds 32[degrees]C, more than 90% of the D. polymorpha in experimental cages died, whereas in the moderately heated zone (maximum summer temperature [less than or equal to] 30[degrees]C) mortality was less than 10% and did not differ from the control, ambient zone (Karatayev 1983). Similar upper maximal max·i·mal
1. Of, relating to, or consisting of a maximum.
2. Being the greatest or highest possible. temperature limits for D. polymorpha survival have been found by other authors in different regions of the FSU: 31.5[degrees]C in Zaporozhskoe Reservoir and 32[degrees]C in a canal of the Pridnieprovskaya Power Station in Ukraine (Lyakhnovich et al. 1994), 32[degrees]C in Kuchurganskiy Liman in Moldova (Vladimirov 1983), 33[degrees]C in a cooling reservoir of the South-Ukrainian Nuclear Power Station (Sinitsina & Protasov 1993) and 34[degrees]C in a cooling reservoir of the Chernobyl Nuclear Power Station (Protasov et al. 1983).
The upper temperature limit for D. polymorpha in European cooling reservoirs is similar to that found in the Lower Mississippi River
River, southern Louisiana, U.S. A distributary of the Red and Mississippi rivers, it branches southwest from the Red River in east-central Louisiana and flows south about 140 mi (225 km) to Atchafalaya Bay. Its name is Choctaw for “Long River.” system (Louisiana) D. polymorpha grow throughout the winter and growth increases in late spring (April and May). Adult mortality occurs from May to August because dissolved oxygen levels decline and minimum daily temperatures warm above 29[degrees]C in the floodplain floodplain, level land along the course of a river formed by the deposition of sediment during periodic floods. Floodplains contain such features as levees, backswamps, delta plains, and oxbow lakes. and 32.5[degrees]C in riverine riv·er·ine
1. Relating to or resembling a river.
2. Located on or inhabiting the banks of a river; riparian: "Members of a riverine tribe ... sites. Thus, the maximum upper temperature limit for D. polymorpha survival is similar in both Europe and North America.
In temperate regions, zebra mussel growth stops in the winter and resumes in the spring after water temperatures warm. Although Smit et al. (1992) assumed that the lower temperature limit for shell growth should be set by the lower temperature limit for filtering (3[degrees]C; Mikheev 1967a, 1967b, Kondratiev 1969, Reeders & bij de Vaate 1990), a majority of studies have found that the threshold temperature for mussel growth is 10[degrees]C to 12[degrees]C (12[degrees]C, Kachanova 1961; 11[degrees]C, Morton 1969a, 1969b; 10[degrees]C, Alimov 1974, Karatayev 1983, Jantz & Neumann 1992; 10[degrees]C to 12[degrees]C, Mackie 1991). bij de Vaate (1991) did report a lower temperature limit of 6[degrees]C, however in North America MacIsaac (1994) found that small mussels incubated at 6[degrees]C experienced shell degrowth and mass loss, whereas large individuals experienced shell degrowth but weight gain. These differences among studies may be the result of local effects, but clearly call for further study.
Maximum growth in D. polymorpha is usually found early in the growing season (Karatayev 1983, 1984, Smit et al. 1992, Lvova et al. 1994, Burlakova 1998, Garton & Johnson 2000), and corresponds with a peak in phytoplankton phytoplankton
Flora of freely floating, often minute organisms that drift with water currents. Like land vegetation, phytoplankton uses carbon dioxide, releases oxygen, and converts minerals to a form animals can use. abundance (Walz 1978a). In midsummer growth rate often decreases (Spiridonov 1971, Walz 1978a, Smit et al. 1992, Stanczykowska & Lewandowski 1995, Allen et al. 1999) and has been attributed to low food concentrations (Walz 1978a), blooms of dinoflagellates dinoflagellates
minute aquatic protozoa; they produce red pigment and toxins which are taken up by shellfish without apparent ill effect, but the toxin is not metabolized and the shellfish may poison animals if eaten. (e.g., Ceratium hirundinella) that impede filter feeding (Stanczykowska & Lewandowski 1995), high water temperatures (Allen et al. 1999) and spawning (Spiridonov 1971, Lvova 1977, 1980, Karatayev 1983, 1992, Allen et al. 1999). In the autumn, when water temperatures decrease, growth stops (Morton 1969a). In the Uchinskoe Reservoir growth stops when temperatures fall to 10[degrees]C (Lvova 1977, 1980) and in the River Rhine at 10[degrees] to 15[degrees]C (Jantz & Neumann 1992).
Based on a bioenergetics bioenergetics,
n 1. system in which natural healing is enhanced by creating harmony between the patient's body and the natural environment.
2. model of zebra mussel growth in the Laurentian Great Lakes Great Lakes, group of five freshwater lakes, central North America, creating a natural border between the United States and Canada and forming the largest body of freshwater in the world, with a combined surface area of c.95,000 sq mi (246,050 sq km). , Schneider (1992) predicted positive growth in the spring and fall when high phytoplankton biomass associated with spring and fall turnover coincides with temperatures near the optimum for growth. Even under conditions of high food availability, growth rates in his model typically decline in the beginning of summer and increase again in August and September as temperature begins to decline. Schneider (1992) used Walz's bioenergetic estimates of metabolic parameters for zebra mussels from Lake Constance Noun 1. Lake Constance - a lake in southeastern Germany on the northern side of the Swiss Alps; forms part of the Rhine River
Deutschland, FRG, Germany, Federal Republic of Germany - a republic in central Europe; split into East Germany (Walz 1978d), where the optimal temperature range for growth is 8[degrees]C to 15[degrees]C. This temperature range is much lower than optimum found by other authors, and therefore more empirical tests of this model are needed before we can assess the generalizability of its predictions.
Location in the Water Column
Dreissena polymorpha grow faster in the water column above the bottom (e.g., on buoys, cages, submerged constructions, floating objects) than on the bottom (Kachanova 1963, Mikheev 1964, bij de Vaate 1991, Smit et al. 1992, 1993, Dorgelo 1993, Burlakova 1998). Yu and Culver (1999) tested the effect of cage location in stratified stratified /strat·i·fied/ (strat´i-fid) formed or arranged in layers.
Arranged in the form of layers or strata. Hargus Lake (Ohio), and found highest growth at their pelagic pelagic
living in the middle or near the surface of large bodies of water such as lakes or oceans. site (2.5-4 m depth) and in the littoral zone littoral zone: see ocean. at 2.5 m depth. All mussels held below the thermocline ther·mo·cline
A layer in a large body of water, such as a lake, that sharply separates regions differing in temperature, so that the temperature gradient across the layer is abrupt. (5-m depth) died before the end of experiment (163 days).
Trophic conditions also affect zebra mussel growth (Table 1). Dorgelo (1993) found that zebra mussels growth rates in Dutch eutrophic lakes A eutrophic lake is a lake with high primary productivity, the result of high nutrient content. These lakes are subject to excessive algal blooms, resulting in murky water and poor water quality. Mararsseveen II and Vechten was higher (0.54-0.59 mm [wk.sup.-1]) than of those grown in mesooligotrophic Lake Masrsseveen I (0.35 mm [wk.sup.-1]), even though there was no difference in mean temperature between these lakes. Jantz and Neumann (1992) found a significant strong correlation ([r.sub.s] = 0.80) between the rate of shell length growth and chlorophyll a Noun 1. chlorophyll a - a blue-black plant pigment having a blue-green alcohol solution; found in all higher plants
chlorophyl, chlorophyll - any of a group of green pigments found in photosynthetic organisms; there are four naturally occurring forms concentration and between shell growth and temperature ([r.sub.s] = 0.82). However, because these two environmental factors are highly correlated, it is impossible to determine the relative contribution of each of these factors on growth (Jantz & Neumann 1992). In a later study (Jantz & Neumann 1998) they found that shell growth rate and the duration of the growing season were correlated with the quantity of available algal algal
pertaining to or caused by algae.
is very rare but systemic and udder infections are recorded. See protothecosis.
the algae Prototheca trispora and P. food.
Sprung (1995a) found a strong correlation between zebra mussel shell growth and food conditions (seston concentration). He suggested that this correlation will exist when seston concentrations stay below those at which the intestine is filled to capacity when the animal filters at a maximum rate (Sprung 1995b). Similarly, Schneider et al. (1998) found that the scope for growth under laboratory conditions had a strong positive relationship with food quality.
In areas with constant water current D. polymorpha grow faster than in still water (Table 1). Kachanova (1963) found that D. polymorpha grow faster on the concrete walls of the canal flowing from Uchinskoe Reservoir than in the reservoir. Mikheev (1964) found that in Kuybyshevskoe Reservoir moderate water currents (up to 0.8 m [s.sup.-1]) facilitate mussel feeding and respiration respiration, process by which an organism exchanges gases with its environment. The term now refers to the overall process by which oxygen is abstracted from air and is transported to the cells for the oxidation of organic molecules while carbon dioxide (CO and D. polymorpha grown in water currents reached 27-28 mm, whereas same aged mussels at the same depth out of currents were only 19-20 mm in length. Smit et al. (1993) suggested that water movement seems to have a larger influence on growth than the amount of algal food in the water column. They found that the young-of-the-year zebra mussels in the Rhine River Rhine River
River, western Europe. Rising in the Swiss Alps, it flows north and west through western Germany to drain through the delta region of The Netherlands into the North Sea. It is 820 mi (1,319 km) long and navigable for 540 mi (870 km). were almost 3 times longer (16 mm) that in Lake Ijsselmeer (6 mm) in spite of lower chlorophyll a concentrations in the river (10-42 [micro]g [L.sup.-1] in the river, 34-106 [micro]g [L.sup.-1] in lakes).
However, strong water currents may inhibit Dreissena growth. The maximum length of 2-y-old zebra mussels in waterways The list of waterways is a link page for any river, canal, estuary or firth.
Electricity produced from generators driven by water turbines that convert the energy in falling or fast-flowing water to mechanical energy. plant with constant water currents <0.5 m [s.sup.-1 was 18 mm, and in places with water currents >1.5 m [s.sup.-1] was 13-14 mm (Mikheev 1964).
Wave action can also inhibit D. polymorpha growth. Mikheev (1964) found that in the littoral zone of Kuybyshevskoe Reservoir exposed to waves, the average (4-5 mm) and maximum (8-10 mm) length of yearling yearling
an animal in its second year of age, e.g. yearling cattle, yearling filly, yearling colt.
rinderpest in wildebeeste in the Serengheti. mussels was almost half that of mussels at the same depth but without waves (7.2 mm mean, 14 mm maximum). He also found that the average length of the young-of-the-year D. polymorpha in parts of the Tsimlyanskoe Reservoir exposed to strong waves was 9 mm (maximum 12.5 mm), whereas in quiet areas at the same depth the average length was 12 mm (maximum 19.2 mm).
Dreissena polymorpha grow faster in shallow than in the deep parts of a waterbody (Table 1). In Kuybyshevskoe Reservoir the maximum length of yearling mussels at 1-1.5 m depth was 13.7-14 mm, and at 20 m depth, only 6-7 mm (Mikheev 1964). Similarly, Garton and Johnson (2000) found that in Lake Wawasee Lake Wawasee, formerly Turkey Lake is a large lake south of Syracuse in Kosciusko County, Indiana, United States. It is the largest natural lake in Indiana. History
Turbidity turbidity /tur·bid·i·ty/ (ter-bid´i-te) cloudiness; disturbance of solids (sediment) in a solution, so that it is not clear.tur´bid
The cloudiness or lack of transparency of a solution.
High concentrations of suspended matter in the water negatively affects filtration, ingestion ingestion /in·ges·tion/ (-chun) the taking of food, drugs, etc., into the body by mouth.
1. The act of taking food and drink into the body by the mouth.
2. , assimilation and growth potential of zebra mussels (Reeders et al. 1989, Noordhuis et al. 1992, Alexander et al. 1994, Summers et al. 1996, Madon et al. 1998, Schneider et al. 1998). In Dutch lakes Dutch Lake is a lake located in the state of Minnesota, the land of 10,000 lakes. It was named for its early German population. It is a 159.50 acre (0 ha) acre lake that is west-southwest of Minneapolis-St. , clearance rates The area which would be cleared per unit time with a stated minimum percentage clearance, using specific minehunting and/or minesweeping procedures. of adult 20-mm zebra mussels declined exponentially as dry suspended matter increased from 5-90 mg [L.sup.-1] (Reeders et al. 1989, Noordhuis et al. 1992).
Madon et al. (1998) found that concentrations of suspended inorganic sediment above 1 mg [L.sup.-1], and a ratio of inorganic to organic fraction of seston higher than 1.71 may cause negative growth. Similar limits were found by Schneider et al. (1998): the scope for growth declined with decreasing food quality and fell below 0 cal [mg.sup.-1] [h.sup.-1] at an organic/inorganic ratio of 0.5. They suggested that high concentrations of suspended inorganic sediment in large turbid tur·bid
Having sediment or foreign particles stirred up or suspended; muddy; cloudy.
tur·bidi·ty n. rivers represents a difficult growth environment for zebra mussels and that populations in turbid rivers may not stabilize at the very high densities typical of lentic Adj. 1. lentic - of or relating to or living in still waters (as lakes or ponds)
lake - a body of (usually fresh) water surrounded by land
lotic - of or relating to or living in actively moving water environments.
The growth rates of D. polymorpha in the same waterbody vary significant among years (Lvova 1980, Dorgelo 1993, Chase & Bailey 1999b). Zebra mussels in the Uchinskoe Reservoir with initial shell lengths of 8 mm grew to 21.2 [+ or -] 0.29 mm by the end of the growing season in 1967, to 19.5 [+ or -] 0.27 mm in 1968, and to 16.8 [+ or -] 0.18 mm in 1969 (Lvova 1980). Dorgelo (1993) found the growth rate of D. polymorpha in lakes Maarsseveen I and II was significantly lower in the summer of 1986 than in 1985. In a study of growth and production of D. polymorpha in lakes St. Clair, Erie and Ontario, Chase and Bailey (1999b) estimated shell production as a part of total production (total production = shell + somatic somatic /so·mat·ic/ (so-mat´ik)
1. pertaining to or characteristic of the soma or body.
2. pertaining to the body wall in contrast to the viscera.
adj. + gamete gamete (găm`ēt): see reproduction. production). They found that the site by year interaction (among 5 populations) explained >80% of the variation; differences between sites in production depended on the year examined. Variation in total production depended on variation in somatic and shell production only, as gamete production was relatively constant among years. Chase and Bailey (1999b) hypothesized that, in response to poor environmental conditions, D. polymorpha shifts the allocation of resources allocation of resources
Apportionment of productive assets among different uses. The issue of resource allocation arises as societies seek to balance limited resources (capital, labour, land) against the various and often unlimited wants of their members. from growth (somatic and shell) to reproduction. As individuals cannot predict how long adverse conditions will persist, investment in growth may be unprofitable (Chase & Bailey 1999b).
Lakes versus Reservoirs
Although environmental factors that affect mussel growth such as temperature, food availability and other conditions can differ greatly among waterbodies and among years within the same waterbody, when we compared size-specific growth rates of zebra mussels among studies that all used a similar method (following caged mussels), some patterns emerged. We compared the data for size specific zebra mussel growth from three studies in reservoirs (two different reservoirs, two different years in one reservoir) and six studies conducted in 5 different lakes (4 in Eastern Europe, 1 in North America). We found that the size-specific growth of mussels in reservoirs was consistently higher than that of mussels grown in lakes (Fig. 2). Surprisingly, given the range of likely conditions among water bodies and among years, growth in reservoirs was very consistent ([R.sup.2] = 0.92). For lakes there was more spread ([R.sup.2] = 0.72), but the patterns and rates were similar among studies. Data for the growth of uncaged mussels in the Svisloch River are much more similar to mussels in reservoirs, whereas those grown in cages were more similar to mussels in lakes (Fig. 1 B).
[FIGURE 2 OMITTED]
Methods to Estimate Longevity
Most of the methods used to estimate longevity of D. polymorpha are similar to those used to estimate growth rate: counting annual rings on shells, analysis of the size-frequency distributions and growth under experimental conditions (Table 2).
Counting Annual Rings on Shell
The maximum longevity reported using this method has decreased through time from 17-19 y (Karpevich 1964) to 4-5 y (Draulans & Wouters 1988). Through time some authors have revised their earlier estimates of zebra mussel longevity. Stanczykowska (1964) initially reported a maximum longevity for D. polymorpha of 10-12 y; 11 y later she revised her estimates from these same data to 5 y (Stanczykowska 1975, 1976b). Kachanova (1963) (this author published later under the name LvovaKachanova and Lvova) reported that the maximum life span of zebra mussels in the Uchinskoe Reservoir was 6-11 y, and later revised this estimate to 4 y (Lvova 1980, Table 2). Although the advantage of this method is that it allows an estimate of the age structure of a population by measuring D. polymorpha at a single point in time, as discussed earlier, it is very difficult to separate annual rings formed during winter from other rings.
Analysis of Size-frequency Distributions
For this method the numbers of peaks on a size-frequency histogram histogram
or bar graph
Graph using vertical or horizontal bars whose lengths indicate quantities. Along with the pie chart, the histogram is the most common format for representing statistical data. are counted, assuming that each peak represents an age class. However, as discussed earlier, age classes may not have distinct sizes, making it difficult to estimate longevity based on size-frequency distributions.
Growth Under Experimental Conditions
Usually authors keep mussels of different initial sizes in cages for a limited period (1-4 y), and then the obtained growth rates are used to estimate the time to reach the maximum size found in the population. However keeping mussels in cages can produce different types of artifacts discussed earlier, which may affect observed growth rates and, therefore, estimates of mussel longevity.
Factors Affecting Longevity
To our knowledge, the first estimates of the longevity of zebra mussels were reported by Karpevich (1952) and Clarke (1952) (Table 2). Karpevich (1952) counted annual rings on shells and estimated zebra mussel longevity in the Volga River Volga River
River, western Russia. Europe's longest river and the principal waterway of western Russia, it rises in the Valdai Hills northwest of Moscow and flows 2,193 mi (3,530 km) southeastward to empty into the Caspian Sea. as 18 y. In contrast, Clarke (1952) using unpublished data from J. Wilhelmi (study site not mentioned) found three peaks in the size-frequency distribution of D. polymorpha and suggested that typical longevity is about three years. Overall, the longevity of D. polymorpha estimated by different authors over the last 50 y varies from 2-19 y. However, the maximum sizes of D. polymorpha reported by these authors are similar (Table 2). This contradiction supports a hypothesis that the reported differences in longevity may be explained to a large extent by the artifacts of the methods used. This suggestion is also supported by the fact that the average D. polymorpha longevity estimated by counting annual rings on shells (7.4 [+ or -] 0.9) is significantly different from average longevity estimated by analysis of size-frequency distribution (3.3 [+ or -] 0.3, P < 0.001, 2-sided t-test) (Table 2). Alternatively, zebra mussels may have a fixed maximum size, and local conditions that affect growth rates determine longevity--fast growing mussels will live for shorter periods of time, whereas slow growing mussels will live longer. Therefore, it is unclear how much of this variability in longevity is natural or is caused by the artifacts of the methods used and definitely requires future investigation.
GENERAL FINDINGS AND FUTURE DIRECTIONS
Although many generalizations can be made about the growth rate and longevity of D. polyrnorpha, and the impacts of various environmental factors on these parameters, the answers to many questions are far from clear. The most important questions that need to be addressed, problems that need to be solved, and targets for future study are:
Growth rate and longevity of D. polymorpha have been estimated by using four different methods, most of which have serious methodological problems. Thus, different estimates of D. polymorpha growth rates and longevity are affected not only by differences in environmental conditions but also by artifacts of the methods used. Following the growth of undisturbed un·dis·turbed
Not disturbed; calm.
1. quiet and peaceful: an undisturbed village
2. D. polymorpha will provide more reliable data on growth potential and variability among different waterbodies with different environmental conditions. Following and subsampling For the signal processing technique, see .
In computer graphics, subsampling (or "downsampling") is the process of reducing an image to a smaller size. It is a type of image scaling, usually used to alter the appearance of an image or reduce the quantity of information required mussels that naturally settle on experimental surfaces through time provides the control and ease found in experimental studies with the growth rates expected from natural populations.
Co-effects of Environmental Factors
Dreissena polymorpha growth rates depend on water temperature, season of the year, location in the water column, food availability, oxygen concentrations, water velocity and various other environmental factors (Table 1). However, it is difficult to separate the independent effects of each of these factors, especially in natural waterbodies where most of these factors will covary. Several factors may have additive or synergistic effects, making it difficult to study the effect of a single factor. Separation of the effects of single and combined factors on growth is essential.
The upper temperature limit for zebra mussel growth seems to be 30[degrees]C to 32[degrees]C, and the lower temperature limit ~10[degrees]C to 12[degrees]C (Kachanova 1961, Morton 1969a, 1969b, Alimov 1974, Karatayev, 1983, Mackie 1991, Jantz & Neumann 1992). However, some studies have found much lower limits (bij de Vaate 1991, Smit et al. 1992). Differences among studies may be a result of local effects but clearly calls for further study.
Growth in Different Types of Waterbodies
There seems to be substantial differences in growth between mussels in reservoirs and lakes--mussels grow much faster in reservoirs than lakes. Experiments that directly test the relative contributions of environmental factors versus the type of water body and what factors are different between reservoirs and lakes are clearly called for to answer this question. It may be that reservoirs provide a better overall growth environment in terms of temperature, nutrition, and water motion than do natural lakes or rivers.
The reported longevity of D. polymorpha varies from 2 to 19 y. It is critically important to understand to what extent this variation is caused by biological variability, environmental conditions and what amount of the variation is caused by the methods used. In addition to the basic value of understanding the variability in D. polymorpha longevity, it is also important if we are to predict population dynamics, spread or to develop control methods for this important invader.
The authors acknowledge the support provided by Stephen F. Austin State University Stephen F. Austin is one of four public universities in Texas not affiliated with a university system. Academics
Stephen F. Austin offers more than 120 areas of study, including more than 80 undergraduate majors, nearly 60 graduate degrees, and two doctoral programs. Stephen F. (Faculty Research Grant # 14123 to AYK AYK As You Know , LEB LEB Liga Española de Baloncesto
LEB London Electricity Board (UK)
LEB Listeria Enrichment Broth
LEB Lebanon/Hanover/White River, NH, USA - Lebanon Regional (Airport Code)
LEB Lower Equipment Bay and DKP DKP Deutsche Kommunistische Partei (German Communist Party)
DKP Diketopiperazine (aspartame by-product)
DKP Dragon Kill Points (massively multiplayer online games) , 2003 to 2004). In the Republic of Belarus the research was supported by a grant from the Ministry of Education and Science Republic of Belarus, grant number 657/65. This work was conducted while D. Padilla was a Sabbatical sab·bat·i·cal also sab·bat·ic
1. Relating to a sabbatical year.
2. Sabbatical also Sabbatic Relating or appropriate to the Sabbath as the day of rest.
A sabbatical year. Fellow at the National Center for Ecological Analysis and Synthesis The National Center for Ecological Analysis and Synthesis is a research center for the science of ecology, located in Santa Barbara, California, USA. Better known by its acronym NCEAS (pronounced N-seece), it opened in May, 1995, funded by the US National Science Foundation, the , a center funded by NSF NSF - National Science Foundation (Grant #DEB-0072909), the University of California The University of California has a combined student body of more than 191,000 students, over 1,340,000 living alumni, and a combined systemwide and campus endowment of just over $7.3 billion (8th largest in the United States). and the Santa Barbara Santa Barbara (săn'tə bär`brə, –bərə), city (1990 pop. 85,571), seat of Santa Barbara co., S Calif., on the Pacific Ocean; inc. 1850. campus.
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1. The collection of organisms living on or in sea or lake bottoms.
2. The bottom of a sea or lake.
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The region of land extending from the backshore to the beginning of the offshore zone.
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Of, relating to, or being the geographic areas adjacent to the Tropics.
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ALEXANDER Y. KARATAYEV, (1) * LYUBOV E. BURLAKOVA (1) AND DIANNA K. PADILLA (2,3)
(1) Department of Biology, Stephen F. Austin State University, Nacogdoches, Texas Nacogdoches (pronounced [ˌnæːkə̆ˈdoʊtʃɪs]) is a city in Nacogdoches County, Texas, in the United States. As of the 2000 census, the city population was 29,914. 75962; (2) Department of Ecology and Evolution, Stony Brook University The State University of New York at Stony Brook (SUNYSB), also known as Stony Brook University (SBU) is a public research university located in Stony Brook, New York (on the north side of Long Island, about 55 miles east of Manhattan, New York). Stony Brook, New York
Stony Brook is a hamlet (unincorporated community) (and census-designated place) located in the Town of Brookhaven in Suffolk County, New York. The population was 13,727 at the 2000 census. 11794; (3) National Center for Ecological Analysis and Synthesis, 735 State Street Suite 300, Santa Barbara, California Santa Barbara is a city in California, United States. It is the county seat of Santa Barbara County, California. As of the 2000 census, the city had a total population of 92,325. 93101
* Corresponding author. E-mail: firstname.lastname@example.org
TABLE 1. Impact of environmental factors on the growth rate of Dreissena polymorpha. Factor Impact Temperature increase Accelerates growth if maximum temperature <30[degrees]C Season of a year Maximum growth is usually at the beginning of the growing season Location in the water column Growth is faster in the water column than on the bottom Trophic conditions Growth is faster in eutrophic than oligotrophic waters Water current Moderate current accelerates growth Depth Growth decreases with depth Wave action Inhibits growth rate Turbidity High amount of suspended matter inhbits growth rate Year-to-year variation Growth varies significantly Factor References Temperature increase Mikheev 1964, Morton 1969a, Spiridonov 1969, Yaroshenko & Naberezhnyi 1971, Lvova-Kachanova 1972, Skalskaya 1976a, 1976b, Stanczykowska 1976a, Kornobis 1977, Elagina et al. 1978, Walz 1978b, 1978c, Karatayev & Tishchikov 1979, Lvova 1980, Karatayev 1983, 1984, 1988, Smit et al. 1992, 1993, Maclsaac 1994, Jantz 1996 Season of a year Mikheev 1964, Lovova 1980, Karatayev 1983, Sprung 1995a, Jantz 1996, Burlakova 1998 Location in the water column Kachanova 1963, Spiridonov 1971, Kornobis 1977, bij de Vaate 1991, Yu & Culver 1999 Trophic conditions Walz 1978a, Dorgelo & Gorter 1984, Smit et al. 1992, 1993, Dorgelo 1993, Sprung 1992, 1995a, Jantz 1996, Burlakova 1998, Jantz & Neumann 1998, Schneider et al. 1998, Horvath & Lamberti 1999 Water current Kachanova 1963, Mikheev 1964, bij de Vaate 1991, Smit et al. 1992, 1993, Dorgelo 1993, Burlakova 1998 Depth Mikheev 1964, Garton & Johnson 2000 Wave action Mikheev 1964 Turbidity Reeders et al. 1989, Noordhuis et al. 1992, Alexander et al. 1994, Summers et al. 1996, Madon et al. 1998, Schneider et al. 1998 Year-to-year variation Lvova 1980, Dorgelo 1993, Chase & Bailey 1999b TABLE 2. Estimates of the longevity of Dreissena polymorpha from different methods. Maximum Waterbody Longevity (years) Length (mm) Counts of annual rings on shells Volga River 18 32-33 Volga River 17-19 30-32 Uchinskoe Reservoir 11 32 Pyalovskoe Reservoir 10-12 29-33 Mazurian lakes 5-7, max 10 n.r. Firth Szczecin 5-6 30-35 Volgogradskoe Reservoir 7-9 30 Masurian lakes 5 n.r. Koninskie lakes 4 29 Lukomskoe Lake 6-8 32-34 Jorzec Lake 7 n.r. Glebokie Lake 5 n.r. Bartag Lake 5 n.r. Otow Lake 5 n.r. Majcz Wielki Lake 5 n.r. Inulec Lake 5 n.r. Zelwazek Lake 5 n.r. Plas Leblance Pond 5 29.9 Laguno Pond 4 25.7 Tsimlyanskoe Reservoir 7 31-33 Average [+ or -] SE 7.4 [+ or -] 0.9 Analysis of size-frequency distributions n.r. 3 31-34 Reservoir #2 Walthamstow 5 40 River Rhine 3 31 Lake Esrom 4 32 River Svisloch 3 30 Lake Ontario: Stoney Point 2 n.r. Whealtey (2 and 6 m) 2-3 n.r. Port Dalhousie (2 and 6 m) [greater than or equal n.r. to] 4 Average [+ or -] SE 3.3 [+ or -] 0.3 Growth in experimental cages Uchinskoe Reservoir 4 36 Lukomskoe Lake 8 30 Lake Wawasee 3 n.r. Svisloch River 3 30 Waterbody Reference Volga River Karpevich 1952 Volga River Karpevich 1964 Uchinskoe Reservoir Kachanova 1963 Pyalovskoe Reservoir Mikheev 1964 Mazurian lakes Stanczykowska 1963 Firth Szczecin Wiktor 1969 Volgogradskoe Reservoir Spiridonov 1971 Masurian lakes Stanczykowska 1975 Koninskie lakes Kornobis 1977 Lukomskoe Lake Karatayev & Tishchikov 1979 Jorzec Lake Lewandowski 1982b Glebokie Lake Lewandowski 1982b Bartag Lake Lewandowski 1982b Otow Lake Lewandowski 1982b Majcz Wielki Lake Stanczykowska et al. 1983 Inulec Lake Stanczykowska et al. 1983 Zelwazek Lake Stanczykowska et al. 1983 Plas Leblance Pond Draulans & Wouters 1988 Laguno Pond Draulans & Wouters 1988 Tsimlyanskoe Reservoir Miroshnichenko 1990 Average [+ or -] SE n.r. Clarke 1952 Reservoir #2 Walthamstow Morton 1969a River Rhine Jantz & Neumann 1992 Lake Esrom Dall & Hamburger 1996 River Svisloch Burlakova 1998 Lake Ontario: Stoney Point Chase & Bailey 1999a Whealtey (2 and 6 m) Chase & Bailey 1999a Port Dalhousie (2 and 6 m) Chase & Bailey 1999a Average [+ or -] SE Uchinskoe Reservoir Lvova 1980 Lukomskoe Lake Karatayev 1984 Lake Wawasee Garton & Johnson 2000 Svisloch River Burlakova 1998 n.r. = not reported