Effect of habitat biogeochemicals on survival of lobsters (Homarus americanus).ABSTRACT Lobsters from eastern Long Island Sound (LIS LIS - Langage Implementation Systeme. A predecessor of Ada developed by Ichbiah in 1973. It was influenced by Pascal's data structures and Sue's control structures. A type declaration can have a low-level implementation specification. ) were exposed to laboratory conditions designed to simulate the habitat in western LIS during the 1999 mass mortality. Elsewhere (Robohm et al. 2005) we have shown that hypoxia hypoxia Condition in which tissues are starved of oxygen. The extreme is anoxia (absence of oxygen). There are four types: hypoxemic, from low blood oxygen content (e.g., in altitude sickness); anemic, from low blood oxygen-carrying capacity (e.g. and concomitant biogeochemical conditions increase the susceptibility of lobsters to a common pathogen at normal summer temperatures. Here we report that in otherwise favorable conditions, high temperature (24[degrees]C) alone had no lethal effect on disease-free eastern LIS lobsters for weeks. However, at this temperature, moderate hypoxia (<2.5 mg/L dissolved oxygen) was lethal to half of a group (n = 20) of lobsters in about 5 days. Addition of sulfide and ammonium to levels reported for LIS (5.5 [micro]M sulfide and 17 [micro]M ammonium) decreased the L[T.sub.50] to 3.3 days. Higher sulfide concentrations decreased the L[T.sub.50] significantly. We conclude that lobster survival times are greatly diminished by hypoxia, sulfide and ammonium at high temperature. Because seasonal hypoxia with the production of sulfide and ammonium is an annual occurrence due to cultural eutrophication Cultural eutrophication is the process that speeds up natural eutrophication because of human activity. Due to clearing of land and building of towns and cities, runoff water is accelerated and more nutrients such as phosphates and nitrate are supplied to the lakes and ponds. in western LIS, the results imply that habitat conditions could have been sufficient to account for the observed mortalities in the warm summer of 1999. KEY WORDS: lobster, Homarus americanus, hypoxia, sulfide, ammonium, temperature INTRODUCTION Lobster mortality has frequently been associated with hypoxia (e.g., Thomas et al. 1979, Hagopian & Riley 1989, Baden et al. 1990, Hallbaeck & Ulmestrand 1990). In 1999, American lobsters (Homarus americanus H. Milne Edwards, 1837) worth perhaps $50 million died in the western one third of the Long Island Sound (LIS). Among the causes offered to explain the event (Pearce & Balcom 2005) were a biologic infection exacerbated by environmental stressors (high temperature, low dissolved oxygen, and high concentrations of sulfide and ammonium). However, the potency and dynamics of this potential mechanism were not understood. Because a parasitic amoeba amoeba: see ameba. amoeba One-celled protozoan that can form temporary extensions of cytoplasm (pseudopodia) in order to move about. Some amoebas are found on the bottom of freshwater streams and ponds. was found in affected lobsters, we initially examined the effect of habitat biogeochemicals and temperature on disease resistance (Robohm et al. 2005). As the meteorologic me·te·or·ol·o·gy n. The science that deals with the phenomena of the atmosphere, especially weather and weather conditions. [French météorologie, from Greek and hydrographic hy·drog·ra·phy n. pl. hy·drog·ra·phies 1. The scientific description and analysis of the physical conditions, boundaries, flow, and related characteristics of the earth's surface waters. 2. data for LIS in 1999 were examined (Wilson & Swanson 2005), it became clear that much higher than normal temperatures had occurred in the bottom water of LIS in 1999. Seasonal bottom water biogeochemical conditions in western LIS arise as a consequence of cultural eutrophication emanating from the New York City New York City: see New York, city. New York City City (pop., 2000: 8,008,278), southeastern New York, at the mouth of the Hudson River. The largest city in the U.S. metropolitan area. The organic matter resulting from the nutrient-stimulated planktonic plank·ton n. The collection of small or microscopic organisms, including algae and protozoans, that float or drift in great numbers in fresh or salt water, especially at or near the surface, and serve as food for fish and other larger organisms. overproduction o·ver·pro·duce tr.v. o·ver·pro·duced, o·ver·pro·duc·ing, o·ver·pro·duc·es To produce in excess of need or demand. o falls out of the photic zone The photic zone or euphotic zone (Greek 'well lit') is the depth of the water whether in a lake or an ocean, that is exposed to sufficient sunlight for photosynthesis to occur. The depth of the euphotic zone can be greatly affected by seasonal turbidity. (Parker & O'Reilly 1991), enriching the near bottom water and sediment and depleting dissolved oxygen. Hypoxia itself limits lobster survival (McLeese 1956, Dias & Rosenberg 1995), but seasonal declines in dissolved oxygen result in secondary changes in the biogeochemistry bi·o·ge·o·chem·is·try n. The study of the relationship between the geochemistry of a region and the animal and plant life in that region. bi of lobster habitat. Under hypoxic hypoxic a state of hypoxia. hypoxic cell sensitizers compounds that selectively sensitize hypoxic tumor cells to the effects of radiation. and anoxic an·ox·i·a n. 1. Absence of oxygen. 2. A pathological deficiency of oxygen, especially hypoxia. [an- + ox(o)- + -ia1. conditions, nitrogen in the organic matter is released as physiologically active ammonium rather than completely oxidized oxidized having been modified by the process of oxidation. oxidized cellulose see absorbable cellulose. nitrate (Froelich et al. 1979). In addition, after oxygen is depleted de·plete tr.v. de·plet·ed, de·plet·ing, de·pletes To decrease the fullness of; use up or empty out. [Latin d , the heterotrophic heterotrophic /het·ero·tro·phic/ (-tro´fik) not self-sustaining; said of microorganisms requiring a reduced form of carbon for energy and synthesis. benthic ben·thos n. 1. The collection of organisms living on or in sea or lake bottoms. 2. The bottom of a sea or lake. [Greek. microbes use a sequence of terminal electron acceptors ([Mn.sup.4+], N[O.sub.3.sup.-], N[O.sub.2.sup.-], [Fe.sup.3+], S[O.sub.4.sup.2-]) to continue oxidization of organic material (Wang & Chapman 1999). Sulfate sulfate, chemical compound containing the sulfate (SO4) radical. Sulfates are salts or esters of sulfuric acid, H2SO4, formed by replacing one or both of the hydrogens with a metal (e.g., sodium) or a radical (e.g., ammonium or ethyl). is notable both because it is abundant in seawater seawater Water that makes up the oceans and seas. Seawater is a complex mixture of 96.5% water, 2.5% salts, and small amounts of other substances. Much of the world's magnesium is recovered from seawater, as are large quantities of bromine. (x100 the dissolved oxygen concentration) and because the reduced product (sulfide) is biologically active. The seasonal process starts in spring in deeper sediment layers with remnant winter conditions. Sediment is anoxic below some horizon where microbial microbial pertaining to or emanating from a microbe. microbial digestion the breakdown of organic material, especially feedstuffs, by microbial organisms. respiration exceeds the oxygen resupply re·sup·ply tr.v. re·sup·plied, re·sup·ply·ing, re·sup·plies To provide with fresh supplies, as of weapons and ammunition. re from overlying overlying suffocation of piglets by the sow. The piglets may be weak from illness or malnutrition, the sow may be clumsy or ill, the pen may be inadequate in size or poorly designed so that piglets cannot escape. water. As the summer progresses, bottom water oxygen concentrations decline and respiration rates increase with increasing temperature. Less dissolved oxygen diffuses or is advected by bioturbation bi·o·tur·ba·tion n. The stirring or mixing of sediment or soil by organisms, especially by burrowing or boring. [German : bio-, bio- + Latin turb into the sediment to meet the increased consumption, resupply is exceeded at a shallower level in the sediment profile, and the sulfide-generating stratum moves upward toward the sediment-water interface (Kristiansen et al. 2002). If microbial metabolic rates become sufficiently elevated and the bottom water sufficiently hypoxic, sulfide may diffuse into the overlying water even though it may still be oxic. Lobsters in habitats enriched with organic matter may then be exposed to hypoxia, sulfide and ammonium simultaneously. The temperature survival range for H. americanus is quite broad, from -1 to 30.5[degrees]C though lobsters from New Hampshire New Hampshire, one of the New England states of the NE United States. It is bordered by Massachusetts (S), Vermont, with the Connecticut R. forming the boundary (W), the Canadian province of Quebec (NW), and Maine and a short strip of the Atlantic Ocean (E). avoided temperatures >19[degrees]C to 23.5[degrees]C (Crossin et al. 1998). They can survive abrupt temperature changes including increases of 16[degrees]C or decreases of 20[degrees]C (Lawton & Lavalli 1995). Cooper and Uzmann (1980) summarized the lower lethal oxygen level from McLeese (1956) to range from 0.2 mg/L at 5[degrees]C to 1.2 mg/L at 25[degrees]C. Resistance of marine invertebrates to sulfide varies widely (e.g., Theede et al. 1969, yon Oertzen & Schlungbaum 1972, Vismann 1996) likely reflecting adaptation to the frequency and intensity of exposure, some even making use of the energy in sulfide via symbionts (e.g., Girguis et al. 2002). The L[T.sub.50] for Norway lobster (Nephrops norvegicus) at high sulfide concentrations (500 [micro]M) under oxic conditions is 22 h (Butterworth et al. 2004). Sulfide in oxic seawater is oxidized rapidly (Gregoire & Lacroix 2001) but Cuomo et al. (2005) found sulfide to about 10 [micro]M and they found ammonium to 16 [micro]M in LIS. The effects on H. americanus of hypoxia, sulfide and ammonium in combination at these levels are not known. This report presents results of exploratory laboratory experiments undertaken to define the response of lobsters to combinations of habitat biogeochemicals at elevated temperature, and to consider this response in relation to the 1999 lobster mortality in LIS. METHODS A specially designed flow-through experimental system (Fig. 1) was constructed (Wieczorek & Draxler 2005) to simulate chemical and physical conditions in the bottom waters of Long Island Sound. The system consisted of four gas-tight, 250-L tanks, each capable of accommodating 26 lobsters in individual refuges and supplied with 0.37 L/rain/animal of conditioned seawater. Seawater for the system was pumped from Sandy Hook Bay, New Jersey through coarse sand, pebble and shells followed by aeration aeration /aer·a·tion/ (ar-a´shun) 1. the exchange of carbon dioxide for oxygen by the blood in the lungs. 2. the charging of a liquid with air or gas. aer·a·tion n. , decantation decantation /de·can·ta·tion/ (de?kan-ta´shun) the pouring of a clear supernatant liquid from a sediment. decantation the pouring of a clear supernatant liquid from a sediment. and passage through three, open-bed biologic filters. The water was then chemically conditioned through countercurrent countercurrent /coun·ter·cur·rent/ (-kur?ent) flowing in an opposite direction. countercurrent flowing in an opposite direction. gas exchange (oxygen and nitrogen) to adjust dissolved oxygen levels. Sodium sulfide and ammonium chloride ammonium chloride (əmō`nēəm klôr`īd), chemical compound, NH4Cl, a white or colorless, odorless, water-soluble, cubic crystalline salt with a biting taste, commonly known as sal ammoniac. were added through a metering pump. Temperature was controlled by pumping water from a reservoir through an in-line chiller chill·er n. 1. One that chills. 2. A frightening story, especially one involving violence, evil, or the supernatural; a thriller. chiller Noun 1. and a heater. [FIGURE 1 OMITTED] Lobsters were purchased (Garbo Lobster, Groton Connecticut) as having been caught in eastern Long Island Sound. In previous experiments on the effect of habitat biogeochemicals on the resistance of lobsters to pathogens (Robohm et al. 2005), animals were gradually acclimated (1[degrees]C per 2 days) to 20[degrees]C and randomly allocated to treatments. We continued this same protocol in these trials both to maintain comparability, and because a rapid 4[degrees] change may more closely approximate conditions in the bottom water of LIS in 1999 (Wilson & Swanson 2005) than would a more gradual change. Also, based on Lawton and Lavalli (1995), the temperature change alone was not expected to be a factor. Experimental conditions (Table 1) included temperatures to 24[degrees]C, dissolved oxygen ranging from 1.7-6.0 mg/L, sulfide to 12 [micro]M, and ammonium added at either 0 or 17 [micro]M. Once levels were set, control of environmental variables included: temperature ([+ or -] 0.5[degrees]C), dissolved oxygen ([+ or -] 0.3 mg/L), sulfide ([+ or -] 1 [micro]M) and ammonium ([+ or -] 3 [micro]M). Seawater was ambient for Sandy Hook Bay water with salinities ranging from 25-27 psu, approximately the same as western LIS. Analytical chemistry methods and system control and operation are detailed in Wieczorek and Draxler (2005). Ten to 20 lobsters that had been fed sparingly and had major chelae banded were individually numbered for identification and randomly allocated to treatments. They were not sorted by sex but were approximately evenly divided between males and females. For practical reasons, these experiments were conducted at unnaturally high lobster densities (6-12 lobsters/[m.sup.2]). However, this was partially compensated for by provision of an excess of well-ventilated individual shelters, high water flow rates to prevent accumulation of signaling (e.g., Bushmann & Atema 1997) and metabolic chemicals, by carefully tracking individual lobster locations within the tanks to discern anomalous results for either down- or upstream individuals, and minimization of the numbers where possible. For example, when we expected survival to be brief, we used fewer excess animals (randomly removed daily) to monitor physiology and microbiology. Lobsters were considered dead when they did not revive when placed in fully oxygenated seawater. Survival curves for each treatment were done by Kaplan-Meier (product limit) estimation using SigmaStat 3.0 (SPSS A statistical package from SPSS, Inc., Chicago (www.spss.com) that runs on PCs, most mainframes and minis and is used extensively in marketing research. It provides over 50 statistical processes, including regression analysis, correlation and analysis of variance. , Inc., Chicago, Illinois) software program. This method calculates median and mean survival times based on deaths and on those periodically removed from the study (censored) for bacterial analysis. The Mann-Whitney rank sum test was applied to evaluate differences in L[T.sub.50]s between treatment and control groups. RESULTS Survival of lobsters in Trial 1 is illustrated in Figure 2. Lobsters in oxic control in this and all high temperature controls in the study survived until trials were terminated (Table 1). The lone death in oxic conditions was one lobster in the first day of Trial 1 in the 20[degrees]C oxic tank, which we included to ensure a relation to previous experiments (Robohm et al. 2005) at that temperature. Under moderately hypoxic conditions (1.7 mg/L DO, Trial 1, Tank 3) at 24[degrees]C, the L[T.sub.50] was 5.1 days. When a small amount of sulfide was present (5.5 [micro]M, Trial 1, Tank 4), the L[T.sub.50] was reduced to 3.3 days. At higher sulfide concentrations and moderate hypoxia (1.6-2.5 mg/L), still lower L[T.sub.50]s were measured. [FIGURE 2 OMITTED] Survival in treatments with moderate hypoxia (1.6-2.5 mg/L), sulfide (5.5-12 [micro]M) and ammonium (~17 [micro]M) designated HSA HSA Health Savings Account (US) HSA Human Serum Albumin HSA Human Services Agency (Nevada) HSA Health Services Agency HSA Health and Safety Authority (Ireland) 1-HSA5 in Table 1, may be compared with normoxic controls to which no sulfide or ammonium were added (C1-C5). The mean L[T.sub.50] for HSAs (1.87 [+ or -] 1.05 day) is significantly different from controls (P < 0.01, Mann-Whitney rank sum test) if the end of the trial is taken as the L[T.sub.50] for controls. This expedient is adopted because no deaths occurred in controls. Even at a dissolved oxygen concentration not considered hypoxic (3.9 mg/L) there is a suggestion (Trial 5, Tank 3) that increased sulfide (to 11 [micro]M) decreases L[T.sub.50] resulting in the lowest survival observed (0.5 day). When sulfide and ammonium were present, all lobsters in all trials died over very short time intervals (Table 1) regardless of oxygen concentration. DISCUSSION We found that high temperature alone had no short-term, lethal effect on disease-free lobsters. However, moderate hypoxia (about 2 mg/L) reduced resistance as measured by L[T.sub.50] of LIS lobsters at 24[degrees]C to about 1 wk. This is generally consistent with the finding (USEPA USEPA United States Environmental Protection Agency 2000) that Homarus larvae Larvae, in Roman religion Larvae: see lemures. were the most sensitive to oxygen concentration of seven crustacean crustacean (krŭstā`shən), primarily aquatic arthropod of the subphylum Crustacea. Most of the 44,000 crustacean species are marine, but there are many freshwater forms. species tested for growth effects and with McLeese (1956) for lobsters from Canadian waters, although a direct comparison is not possible as he determined 48-h L[C.sub.50]s. Sulfide at environmentally relevant low levels (about 10 [micro]M or less) combined with hypoxia and ammonium at high temperature shortened the survival time to a few days. The regression of survival on sulfide concentration (Fig. 3) is significant though the relationship is likely nonlinear rather than linear as suggested by the statistical model. High resistance to sulfide toxicity among benthic invertebrates seems to be a characteristic of bivalves and, to a lesser extent, "opportunistic" polychaetes. As in other decapod decapod (dĕk`əpŏd') (Gr.,=10 feet), name for invertebrate animals of the crustacean order Decapoda (phylum Arthropoda) including the crabs, the lobsters and crayfish, and the true shrimps, all having five pairs of legs. crustaceans such as Crangon crangon (Theede et al. 1969, Vismann 1996), our results show that for H. americanus sulfide resistance is low, particularly in moderate hypoxia. Further, an induction period induction period n. The interval between an initial injection of an antigen and the appearance of demonstrable antibodies in the blood. consistently preceded precipitous mortality and resulted in all lobsters in all sulfidic trials dying by the end of the same 24-h interval containing the L[T.sub.50] for that group. [FIGURE 3 OMITTED] McLeese (1956) found that lobsters acclimated to lower oxygen concentrations were better able to tolerate hypoxia than those acclimated to higher concentrations. Because acclimation acclimation /ac·cli·ma·tion/ (ak?li-ma´shun) the process of becoming accustomed to a new environment. ac·cli·ma·tion n. 1. in our experiments was always near oxygen saturation oxygen saturation sO2 The O2 concentration of blood expressed as a ratio of its total O2-carrying capacity; the OS is a measure of the utilization of O2 transport capacity; sO2 at 20[degrees]C (about 6 mg/L), we would expect to have obtained longer L[T.sub.50] values had our lobsters been acclimated to a lower concentration (e.g., 3 mg/ L). If this acclimation effect extends to hypoxia-sulfide exposure and if dissolved oxygen declined gradually permitting some acclimation, LIS lobsters in 1999 might have survived slightly longer than the 1-2 days suggested by our laboratory experiments. We conclude that whereas high temperatures may be lethal to lobsters over time, these laboratory experiments show that survival is greatly diminished by hypoxia and the presence of sulfide and ammonium. Since these habitat chemical conditions are generated annually in western LIS due to cultural eutrophication, they could have been sufficient to account for the observed mortalities in the warm summer of 1999. For the lobster fishery, the results of these laboratory experiments suggest that lobster losses may be minimized by limiting soak times of traps to no more than 2 days when bottom temperatures are abnormally elevated above usual summer time highs. LITERATURE CITED Baden, S. P., L. Phil & R. Rosenberg. 1990. Effects of oxygen depletion on the ecology, blood physiology and fishery of the Norway lobster Nephrops norvegicus. Mar. Ecol. Prog. Ser. 67:141-155. Bushmann, P. J. & J. Atema. 1997. Shelter sharing and chemical courtship signals in the lobster Homarus americanus. Can. J. Fish. Aqua. Sci 54:647-654. Butterworth, K. G., M. K. Grieshaber & A. C. Taylor. 2004. Behavioral and physiological responses of the Norway lobster, Nephrops norvegicus (Crustacea: Decapoda), to sulphide exposure. Mar. Biol. 144:1087-1095. Cooper, R. A. & J. Uzmann. 1980. Ecology of juvenile and adult Homarus americanus. In: J. S. Cobb & B. F. Phillips, editors. The biology and management of lobsters, vol. II. New York New York, state, United States New York, Middle Atlantic state of the United States. It is bordered by Vermont, Massachusetts, Connecticut, and the Atlantic Ocean (E), New Jersey and Pennsylvania (S), Lakes Erie and Ontario and the Canadian province of : Academic Press. pp. 97-142. Crossin, G. T., S. A. Al-Ayoub, S. H. Jury, H. Howell & W. H. Watson III. 1998. Behavioral thermoregulation Thermoregulation The processes by which many animals actively maintain the temperature of part or all of their body within a specified range in order to stabilize or optimize temperature-sensitive physiological processes. in the American lobster Homarus americanus. J. Exp. Biol. 201:365-374. Cuomo, C., R. Valente & D. Dogru. 2005. Seasonal variations in sediment and bottom water chemistry of western Long Island Sound: implications for lobster mortality. J. Shellfish Res. 24(3):805-814. Dias, R. J. & R. Rosenberg. 1995. Marine benthic hypoxia: A review of its ecological effects and the behavioral responses of benthic macrofauna. Oceanogr. Mar. Bio. Ann. Rev. 33:245-303. Froelich, P. N., G. P. Klinkhammer, M. L. Bender, N. A. Luedtke, G. R. Heath, D. Hammond, B. Hartman & V. Maynard. 1979. Early oxidation of organic matter in pelagic sediments of the eastern equatorial Atlantic: suboxic diagenesis diagenesis Sum of all processes, chiefly chemical, that produce changes in a sediment after its deposition but before its final lithification. Usually, not all the minerals in a sediment are in chemical equilibrium, so changes in interstitial water composition or in . Geochim. Cosmochim. Acta. 43:1075-1090. Girguis, P. R., J. J. Childress, J. K. Freytag, K. Klose & R. Stuber. 2002. Effects of metabolite metabolite, organic compound that is a starting material in, an intermediate in, or an end product of metabolism. Starting materials are substances, usually small and of simple structure, absorbed by the organism as food. uptake on proton-equivalent elimination by two species of deep-sea vestimentiferan vestimentiferan n. Any of various marine tubeworms, including the giant tubeworms, of the phylum Vestimentifera, that lack a digestive system, depend on chemoautotrophic bacteria for nutrients, and inhabit areas close to deep sea hydrothermal vents or tubeworm tube·worm n. Any of various chiefly marine worms or wormlike invertebrates of the phyla Annelida, Pogonophora, Phoronida, or Vestimentifera, living within tubular cases made of mineral or chitinous secretions or of aggregated grit. , Riftia pachyptila and Lamellibrachia cf luymesi: proton elimination is a necessary adaptation to sulfide-oxidizing chemoautotrophic chemoautotrophic /che·mo·au·to·tro·phic/ (-aw?to-tro´fik) capable of synthesizing cell constituents from carbon dioxide with energy from inorganic reactions. symbionts. J. Exp. Biol. 205: 3055-3066. Gregoire, M. & G. Lacroix. 2001. Study of the oxygen budget of the Black Sea waters using a 3D coupled hydrodynamical-biogeochemical model. J. Mar. Syst. 31:175-202. Hagopian, D. S. & J. G. Riley. 1989. Aeration of lobster pounds. J. Shellfish Res. 8:480. Hallbaeck, H. & M. Ulmestrand. 1990. Norway lobster in the Kattegat. Fauna Flora (Stockh.) 85:186-192. Kristiansen, K. D., E. Kristensen & E. M. Jensen. 2002. The influence of water column hypoxia on the behaviour of manganese and iron in sandy coastal marine sediment. Estuar. Coast. Shelf Sci. 55:645-654. Lawton, P. & K.L. Lavalli. 1995. Postlarval, juvenile, adolescent, and adult ecology. In: J. R. Factor, editor. Biology of the lobster Homarus americanus. San Diego: Academic Press. pp. 47-88. McLeese, D. W. 1956. Effects of temperature, salinity and oxygen on the survival of the American lobster. J. Fish. Res. Bd. Canada. 13:247-272. Parker, C. & J. E. O'Reilly. 1991. Oxygen depletions in Long Island Sound: an historical perspective. Estuaries 14:248-265. Pearce, J. & N. Balcom. 2005. The 1999 Long Island Sound Lobster Mortality event: findings of the comprehensive research initiative. J. Shellfish Res. 24(3):691-697. Robohm, R. A., A. F. J. Draxler, D. Wieczorek, D. Kapareiko & S. Pitchford. 2005. Effects of environmental stressors on disease susceptibility in American lobsters: a controlled laboratory study. J. Shellfish Res. 24(3):773-779. Theede, H., A. Ponat, H. Hiroki & C. Schlieper. 1969. Studies on the resistance of marine bottom invertebrates to oxygen-deficiency and hydrogen sulfide hydrogen sulfide, chemical compound, H2S, a colorless, extremely poisonous gas that has a very disagreeable odor, much like that of rotten eggs. It is slightly soluble in water and is soluble in carbon disulfide. . Mar. Biol. 2:325-337. Thomas, J. P., J. E. O'Reilly, A. F. J. Draxler, J. Babinchak, C. Robertson, W. C. Phoel, R. I. Waldhauer, C. Evans, A. Matte, M. Cohn, M. Nitkowski & S. Dudley. 1979. Biological Processes: productivity and respiration. In: R.L. Swanson & C. J. Sinderman, editors. Oxygen depletion and associated benthic moralities in New York Bight The New York Bight is a large gulf on the Atlantic Ocean along the coast of North America in the northeastern United States. It is formed by the coastal identation between New Jersey and Long Island around the mouth of the Hudson River at New York Harbor. , 1976. NOAA NOAA abbr. National Oceanic and Atmospheric Administration Noun 1. NOAA - an agency in the Department of Commerce that maps the oceans and conserves their living resources; predicts changes to the earth's environment; professional paper 11. Washington, DC: NOAA. pp. 231-261. USEPA. 2000. Ambient aquatic life water quality criteria for dissolved oxygen (saltwater): Cape Cod to Cape Hatteras. EPA-822-R-00-012. Narragansett, RI: United States Environmental Protection Agency "EPA" redirects here. For other uses see EPA (disambiguation) and Environmental Protection Agency. The Environmental Protection Agency (EPA or sometimes USEPA . Vismann, B. 1996. Sulfide species and total sulfide toxicity in the shrimp Crangon crangon. J. Exp. Mar. Biol. Ecol. 204:141-154. von Oertzen, J. A. & G. Schlungbaum. 1972. Experimentell-okologische Untersuchungen uber 02-Mangel-und [H.sub.2]S Resistenz an marinen Evertebraten der westlichen Ostsee. Beitrage zur Meereskunde 29:79-91. Wang, F. & P. M. Chapman. 1999. Biological implications of sulfide in sediment--A review focusing on sediment toxicity. Envir. Tox. Chem. 18:2526-2532. Wieczorek, D. & A. F. J. Draxler. 2005. A method for exposing lobsters to simulated habitat biogeochemicals and temperatures. J. Shellfish Res. 24(3):767-771. Wilson, R. & R. L. Swanson. 2005. A perspective on bottom water temperature anomalies in Long Island Sound during the 1999 lobster mortality event. J. Shellfish Res. 24(3):825-830. ANDREW F. J. DRAXLER, (1) * RICHARD A. ROBOHM, (2) DANIEL WIECZOREK, (1) DIANE KAPAREIKO (2) AND STEVEN PITCHFORD (2) (1) NOAA-Fisheries, James J. Howard This article is about the politician. For other people similarly named, see James Howard. James John Howard (July 24, 1927 in Irvington, New Jersey – March 25, 1988 in Washington, D.C. Marine Sciences Laboratory, Highlands New Jersey 07732; (2) NOAA-Fisheries, Milford Laboratory, Milford Connecticut 06460 * Corresponding author. E-mail: andrew.draxler@noaa.gov
TABLE 1.
Survival time (L[T.sub.50]) of lobsters from eastern Long Island Sound
at conditions that simulate the biogeochemistry of western LIS
habitats.
Reference conditions are indicated as replicates C1-C5 and tanks with
similar hypoxia, sulfide and ammonium are indicated as HSA1-HSA5.
The mean L[T.sub.50]) (1.87 [+ or -] 1.05 d) for HSAs is significantly
different from the controls (P < 0.01, Mann-Whitney Rank Sum Test)
if the end of the trial time is taken as the LT50s for controls.
Number of
Lobsters Temp DO Sulfide
Trial Tank Rep. at Start ([degrees]C) (mg/L) ([micro]M)
1 1 C1 20 24 5.5
1 2 20 20 5.5
1 3 20 24 1.7
1 4 HSA1 20 24 1.7 5.5
2 1 10 24 2.5
2 2 C2 10 24 5.5
2 3 C3 10 24 5.5
2 4 HSA2 10 24 2.5 7.5
3 1 C4 10 24 5.5
3 2 14 24 5.5 10
3 3 HSA3 10 24 1.6 12
4 1 C5 10 24 6.0
4 2 HSA4 10 24 1.8 6
4 3 HSA5 12 24 1.8 6
5 1 10 24 3.4
5 2 10 24 3.4
5 3 10 24 3.9 8
5 4 10 24 3.9 11
Amm Time to 100% [LT.sub.50]
Trial ([micro]M) Mortality (d) (d)
1 * >14
1 ** >14
1 *** 5.1
1 17 4 3.3
2 * >11
2 * >11
2 * >11
2 17 3 1.8
3 * >8
3 17 2 1.5
3 17 2 1.5
4 * >8
4 17 3 1.4
4 17 3 1.3
5 * >2
5 * >2
5 17 *** 1.4
5 17 2 0.5
* = 100% survival to termination of trial.
** = 95% of lobsters remaining alive at termination of trial.
*** = 10% of lobsters remaining alive at termination of trial.
Rep., replicate designation; Temp., temperature; Amm, ammonium
concentration added. Entries for sulfide and ammonium are blank
when none was added.
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