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    INTERNATIONAL PROGRAMME ON CHEMICAL SAFETY








    ENVIRONMENTAL HEALTH CRITERIA 86





    Mercury - Environmental Aspects








    This report contains the collective views of an international group of
    experts and does not necessarily represent the decisions or the stated
    policy of the United Nations Environment Programme, the International
    Labour Organisation, or the World Health Organization.

    First draft prepared at the National Institute of Health Sciences,
    Tokyo, Japan, and the Institute of Terrestrial Ecology, Monk's Wood,
    United Kingdom


    Published under the joint sponsorship of the United Nations
    Environment Programme, the International Labour Organisation, and the
    World Health Organization


    World Health Organization
    Geneva, 1989

         The International Programme on Chemical Safety (IPCS) is a joint
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    CONTENTS

    ENVIRONMENTAL HEALTH CRITERIA FOR MERCURY - ENVIRONMENTAL ASPECTS

    1. SUMMARY AND CONCLUSIONS

         1.1. Physical and chemical properties
         1.2. Sources in the environment
         1.3. Uptake, elimination, and accumulation in organisms
         1.4. Toxicity to microorganisms
         1.5. Toxicity to aquatic organisms
         1.6. Toxicity to terrestrial organisms
         1.7. Effects of mercury in the field

    2. PHYSICAL AND CHEMICAL PROPERTIES

    3. SOURCES OF MERCURY IN THE ENVIRONMENT

         3.1. Natural and anthropogenic sources and cycling
         3.2. Speciation
         3.3. Levels in the environment
         3.4. Methylation of mercury

    4. UPTAKE, LOSS, AND ACCUMULATION IN ORGANISMS

         4.1. Speciation of mercury
         4.2. Uptake and loss in aquatic organisms
              4.2.1. Microorganisms, plants, and invertebrates
              4.2.2. Fish
                     4.2.2.1  Effects of environmental variables on 
                              uptake by fish
              4.2.3. Studies on more than one type of organism
         4.3. Uptake and loss in terrestrial organisms
         4.4. Accumulation in the field
              4.4.1. General exposure
              4.4.2. Mercury manufacturing and general industrial areas
              4.4.3. Mining activity
              4.4.4. Chloralkali plants
              4.4.5. Mercurial fungicides

    5. TOXICITY TO MICROORGANISMS

         5.1. Toxicity of inorganic mercury
              5.1.1. Single species cultures
              5.1.2. Mixed cultures and communities
         5.2. Toxicity of organic mercury

    6. TOXICITY TO AQUATIC ORGANISMS

         6.1. Toxicity to aquatic plants
         6.2. Toxicity to aquatic invertebrates

              6.2.1. Acute and short-term toxicity to
                     invertebrates
              6.2.2. Behavioural effects
         6.3. Toxicity to fish
              6.3.1. Acute and short-term toxicity to fish
              6.3.2. Reproductive effects and effects on
                     early life stages
              6.3.3. Behavioural effects
              6.3.4. Physiological and biochemical effects
         6.4. Toxicity to amphibia
         6.5. Toxicity to aquatic mammals

    7. TOXICITY TO TERRESTRIAL ORGANISMS

         7.1. Toxicity to terrestrial plants
         7.2. Toxicity to terrestrial animals
              7.2.1. Toxicity to terrestrial invertebrates
              7.2.2. Effects of mercury on birds
                     7.2.2.1  Inorganic and metallic mercury
                     7.2.2.2  Effect of organic mercury on birds
              7.2.3. Effects of mercury on non-laboratory mammals

    8. EFFECTS OF MERCURY IN THE FIELD

    9. EVALUATION

         9.1. The marine environment
         9.2. The freshwater environment
         9.3. The terrestrial environment

    REFERENCES
    

    WHO TASK GROUP ON ENVIRONMENTAL HEALTH CRITERIA FOR MERCURY -
    ENVIRONMENTAL ASPECTS

     Participants

    Dr L.A. Albert, Director, Environmental Pollution Programme, National
         Institute for Research on Biotic Resources, Xalapa, Mexico

    Professor T.W. Clarkson, Division of Toxicology, The University of
         Rochester, School of Medicine and Dentistry, Rochester, USA
          (Chairman)

    Dr R. Elias, Environmental Criteria and Assessment Office, US
         Environmental Protection Agency, Research Triangle Park, North
         Carolina, USA

    Dr J.H.M. Temmink, Department of Toxicology, Agricultural University,
         Biotechnion, Wageningen, Netherlands

    Dr G. Roderer, Fraunhofer Institute for Environmental Chemistry and
         Ecotoxicology, Schmallenberg-Grafschaft, Federal Republic of
         Germany

    Dr R. Koch, Division of Toxicology, Research Institute for Hygiene and
         Microbiology, Bad Elster, German Democratic Republic

    Professor Y. Kodama, Department of Environmental Health, University of
         Occupational and Environmental Health, Kitakyushu, Japan

    Professor P.N. Viswanathan, Ecotoxicology Section, Industrial
         Toxicology Research Centre, Lucknow, India

     Observers

    Mr D.J.A. Davies, Department of the Environment, London, United
         Kingdom

    Dr I. Newton, The Institute of Terrestrial Ecology, Monks Wood
         Experimental Station, Huntingdon, United Kingdom

     Secretariat

    Dr S. Dobson, The Institute of Terrestrial Ecology, Monks Wood
         Experimental Station, Huntingdon, United Kingdom  (Rapporteur)

    Dr M. Gilbert, International Programme on Chemical Safety, World
         Health Organization, Geneva, Switzerland  (Secretary)

    Mr P.D. Howe, The Institute of Terrestrial Ecology, Monks Wood
         Experimental Station, Huntingdon, United Kingdom

    NOTE TO READERS OF THE CRITERIA DOCUMENTS

         Every effort has been made to present information in the criteria
    documents as accurately as possible without unduly delaying their
    publication. In the interest of all users of the environmental health
    criteria documents, readers are kindly requested to communicate any
    errors that may have occurred to the Manager of the International
    Programme on Chemical Safety, World Health Organization, Geneva,
    Switzerland, in order that they may be included in corrigenda, which
    will appear in subsequent volumes.

                               *   *   *

         A detailed data profile and a legal file can be obtained from the
    International Register of Potentially Toxic Chemicals, Palais des
    Nations, 1211 Geneva 10, Switzerland (Telephone no. 988400 - 985850).

    ENVIRONMENTAL HEALTH CRITERIA FOR MERCURY - ENVIRONMENTAL ASPECTS

         A WHO Task Group on Environmental Health Criteria for Mercury -
    Environmental Aspects met at the Institute of Terrestrial Ecology,
    Monks Wood, UK, from 7 to 11 December 1987. Dr B.N.K. Davis welcomed
    the participants on behalf of the host Institution, and Dr M. Gilbert
    opened the meeting on behalf of the three co-sponsoring organizations
    of the IPCS (ILO/UNEP/WHO). The Task Group reviewed and revised the
    draft criteria document and made an evaluation of the risks for the
    environment from exposure to mercury.

         The first draft of this document was prepared by Dr S. Dobson and
    Mr P.D. Howe, Institute of Terrestrial Ecology. Dr M. Gilbert and
    Dr P.G. Jenkins, both members of the IPCS Central Unit, were
    responsible for the overall scientific content and editing,
    respectively.

                                  *  *  *

         Partial financial support for the publication of this criteria
    document was kindly provided by the United States Department of Health
    and Human Services, through a contract from the National Institute of
    Environmental Health Sciences, Research Triangle Park, North Carolina,
    USA - a WHO Collaborating Centre for Environmental Health Effects.

    INTRODUCTION

         There is a fundamental difference in approach between the
    toxicologist and the ecotoxicologist concerning the appraisal of the
    potential threat posed by chemicals. The toxicologist, because his
    concern is with human health and welfare, is preoccupied with any
    adverse effects on individuals, whether or not they have ultimate
    effects on performance or survival. The ecotoxicologist, in contrast,
    is concerned primarily with the maintenance of population levels of
    organisms in the environment. In toxicity tests, he is interested in
    effects on the performance of individuals - in their reproduction and
    survival - only insofar as these might ultimately affect the
    population size. To him, minor biochemical and physiological effects
    of toxicants are irrelevant if they do not, in turn, affect
    reproduction, growth, or survival.

         It is the aim of this document to take the ecotoxicologist's
    point of view and consider effects on populations of organisms in the
    environment. No attempt has been made to link the conclusions reached
    in this document with possible effects on human health. This will only
    be feasible when Environmental Health Criteria 1: Mercury (WHO, 1976),
    which considered the effects of mercury on human health, has been
    updated. Due attention has been given to the persistence in the
    environment and the bioaccumulation and transport of mercury in
    aquatic food chains. These will have implications for human
    consumption of the metal.

         This document, although based on a thorough survey of the
    literature, is not intended to be exhaustive in the material included.
    In order to keep the document concise, only those data which were
    considered to be essential in the evaluation of the risk posed by
    mercury to the environment have been included. Concentration figures
    for mercury in the environment, or in particular species of organism,
    have not been included unless they illustrate specific toxicological
    points. "Snap shot" concentration data, where a causal relationship
    between the presence of the metal and an observed effect is not
    clearly demonstrated, have been excluded.

         The term bioaccumulation indicates that organisms take-up
    chemicals to a greater concentration than that found in their
    environment or their food. 'Bioconcentration factor' is a quantitative
    way of expressing bioaccumulation: the ratio of the concentration of
    the chemical in the organism to the concentration of the chemical in
    the environment or food. Biomagnification refers, in this document, to
    the progressive accumulation of chemicals along a food chain.

    1.  SUMMARY AND CONCLUSIONS

    1.1  Physical and chemical properties

         Mercury is a metal which is liquid at normal temperatures and
    pressures. It forms salts in two ionic states mercury(I) and
    mercury(II). Mercury(II), or mercuric, salts are very much more common
    than mercury(I) salts, and hence it is mercuric salts which will be
    mainly considered here. Mercury also forms organometallic compounds,
    some of which have found industrial and agricultural use.
    "Organometallic" is used here to indicate a covalently-bonded
    compound, and does not include mercury bound to proteins nor salts
    formed with organic acids. These organometallic compounds are stable,
    though some are readily broken down by living organisms, while others
    are not readily biodegraded. Elemental mercury gives rise to a vapour
    which dissolves only slightly in water.

    1.2  Sources in the Environment

         Natural mercury arises from the degassing of the earth's crust
    through volcanic gases and, probably, by evaporation from the oceans.
    Local levels in water derived from mercury ores may also be high
    (up to 80 µg/litre). Atmospheric pollution from industrial production
    is probably low, but pollution of water by mine tailings is
    significant. The burning of fossil fuels is a source of mercury. The
    chloralkali industry and, previously, the wood pulping industry also
    released significant amounts of mercury. Although the use of mercury
    is reducing, high concentrations of the metal are still present in
    sediments associated with the industrial applications of mercury. Some
    mercury compounds have been used in agriculture, principally as
    fungicides.

    1.3  Uptake, Elimination, and Accumulation in Organisms

         Mercuric salts, and, to a much greater extent, organic mercury,
    are readily taken up by organisms in water. Aquatic invertebrates, and
    most particularly aquatic insects, accumulate mercury to high
    concentrations. Fish also take up the metal and retain it in tissues,
    principally as methylmercury, although most of the environmental
    mercury to which they are exposed is inorganic. The source of the
    methylation is uncertain, but there is strong indication that
    bacterial action leads to methylation in aquatic systems.
    Environmental levels of methylmercury depend upon the balance between
    bacterial methylation and demethylation. The indications are that
    methylmercury in fish arises from this bacterial methylation of
    inorganic mercury, either in the environment or in bacteria associated
    with fish gills, surface, or gut. There is little indication that fish
    themselves either methylate or demethylate mercury. Elimination of
    methylmercury is slow from fish (with half times in the order of
    months or years) and from other aquatic organisms. Loss of inorganic
    mercury is more rapid and so most of the mercury in fish is retained

    in the form of methylmercury. Terrestrial organisms are also
    contaminated by mercury, with birds being the best studied. Sea birds
    and those feeding in estuaries are most contaminated. The form of
    retained mercury in birds is more variable and depends on species,
    organ, and geographical site.

    1.4  Toxicity to Microorganisms

         The metal is toxic to microorganisms. Inorganic mercury has been
    reported to have effects at concentrations of the metal in the culture
    medium of 5 µg/litre, and organomercury compounds at concentrations at
    least 10 times lower than this. Organomercury compounds have been used
    as fungicides. One factor affecting the toxicity of the organometal is
    the rate of uptake of the metal by cells. Mercury is bound to the cell
    walls or cell membranes of microorganisms, apparently to a limited
    number of binding sites. This means that effects are related to cell
    density as well as to the concentration of mercury in the substrate.
    These effects are often irreversible, and mercury at low
    concentrations represents a major hazard to microorganisms.

    1.5  Toxicity to Aquatic Organisms

         The organic forms of mercury are generally more toxic to aquatic
    organisms than the inorganic forms. Aquatic plants are affected by
    mercury in the water at concentrations approaching 1 mg/litre for
    inorganic mercury but at much lower concentrations of organic mercury.
    Aquatic invertebrates vary greatly in their susceptibility to mercury.
    Generally, larval stages are more sensitive than adults. The 96-h
    LC50s vary between 33 and 400 µg/litre for freshwater fish and are
    higher for sea-water fish. However, organic mercury compounds are more
    toxic. Toxicity is affected by temperature, salinity, dissolved
    oxygen, and water hardness. A wide variety of physiological and
    biochemical abnormalities has been reported after fish have been
    exposed to sublethal concentrations of mercury, although the
    environmental significance of these effects is difficult to assess.
    Reproduction is also affected adversely by mercury.

    1.6  Toxicity to Terrestrial Organisms

         Plants are generally insensitive to the toxic effects of mercury
    compounds. Birds fed inorganic mercury show a reduction in food intake
    and consequent poor growth. Other, more subtle, effects on enzyme
    systems, cardiovascular function, blood parameters, the immune
    response, kidney function and structure, and behaviour have been
    reported. Organomercury compounds are more toxic for birds than are
    inorganic.

    1.7  Effects of Mercury in the Field

         Pollution of the sea with organomercury led to the death of fish
    and fish-eating birds in Japan. Except for this incident at Minamata,
    few follow-up studies of the effects of localised release have been
    conducted. The use of organomercury fungicides as seed dressings in
    Europe led to the deaths of large numbers of granivorous birds,
    together with birds of prey feeding on the corpses. Residues of
    mercury in birds' eggs have been associated with deaths of embryos in
    shell. The presence of organochlorine residues in the same birds and
    their eggs makes an accurate assessment of the effects of mercury
    difficult. It is, however, thought to be a contributing factor in the
    population decline of some species of raptors.

    2.  PHYSICAL AND CHEMICAL PROPERTIES

         The physical and chemical properties of mercury have been
    detailed in Environmental Health Criteria 1: Mercury (WHO, 1976). The
    relevant chapter is summarized here.

         Mercury can exist in a wide variety of physical and chemical
    states. The different chemical and physical forms of this element all
    have their intrinsic toxic properties and different applications in
    industry and agriculture, and require a separate assessment of risk.

         Mercury, along with cadmium and zinc, falls into Group IIb of the
    Periodic Table. In addition to its elemental state, mercury exists in
    the mercury (I) and mercury (II) states in which the mercury atom has
    lost one and two electrons, respectively. The chemical compounds of
    mercury (II) are much more numerous than those of mercury (I).

         In addition to simple salts, such as chloride, nitrate and
    sulfate, mercury (II) forms an important class of organometallic
    compounds. These are characterized by the attachment of mercury to
    either one or two carbon atoms to form compounds of the type RHgX and
    RHgR' where are R and R' represent the organic moiety. The most
    numerous are those of the type RHgX. X may be one of a variety of
    anions. The carbon-mercury bond is chemically stable. It is not split
    in water nor by weak acids or bases. The stability is not due to the
    high strength of the carbon-mercury bond but to the very low affinity
    of mercury for oxygen. The organic moiety, R, takes a variety of
    forms, some of the most common being the alkyl, the phenyl, and the
    methoxyethyl radicals. If the anion X is nitrate or sulfate, the
    compound tends to be "salt-like" having appreciable solubility in
    water; however, the chlorides are covalent, non-polar compounds that
    are more soluble in organic solvents than in water. From the
    toxicological standpoint, the most important of these organometallic
    compounds is the subclass of short-chain alkyl mercurials in which
    mercury is attached to the carbon atom of a methyl, ethyl, or propyl
    group.

    3.  SOURCES OF MERCURY IN THE ENVIRONMENT

         The sources of mercury have been detailed in Environmental Health
    Criteria 1: Mercury (WHO, 1976). Relevant data are summarized here.

    3.1  Natural and Anthropogenic Sources and Cycling

         The major source of mercury is the natural degassing of the
    earth's crust and amounts to between 25 000 and 125 000 tonnes per
    year. Anthropogenic sources are probably less than natural sources.
    World production of mercury by mining and smelting was estimated at
    10 000 tonnes per year in 1973 and has been increasing at an annual
    rate of about 2%. The chloralkali, electrical equipment, and paint
    industries are the largest consumers of mercury, accounting for about
    55% of the total consumption. Mercury has a wide variety of other uses
    in industry, agriculture, military applications, medicine, and
    dentistry.

         Several of man's activities, not directly related to mercury,
    account for substantial releases into the environment. These include
    the burning of fossil fuel, the production of steel, cement, and
    phosphate, and the smelting of metals from their sulfide ores.

         Alkylmercury fungicides used as seed dressings are important
    original sources of mercury in terrestrial food chains, although the
    use of these materials has decreased considerably.

         Two cycles are believed to be involved in the environmental
    transport and distribution of mercury. One is global in scope and
    involves the atmospheric circulation of elemental mercury vapour from
    sources on land to the oceans. However, the mercury content of the
    oceans is so large, at least 70 million tonnes, that the yearly
    increases in concentration due to deposition from the global cycle are
    not detectable.

         The other cycle is local in scope and depends upon the
    methylation of inorganic mercury mainly from anthropogenic sources.
    Many steps in this cycle are still poorly understood, but it is
    believed to involve the atmospheric circulation of dimethylmercury
    formed by bacterial action.

    3.2  Speciation

         The following speciation among mercury compounds has been
    proposed by Lindquist et al. (1984), where V stands for volatile, R
    for water-soluble or particle-borne reactive species, and NR for non-
    reactive species (Hg° is elemental mercury):

         V:   Hg°, (CH3)2Hg

         R:   Hg2+, HgX2, HgX3-, and HgX42-,

              with X = OH-, Cl- and Br-.

              HgO on aerosol particles. Hg2+ complexes with organic
              acids.

         NR:  CH3Hg+, CH3HgCl, CH3HgOH and other organomercuric
              compounds, Hg(CN)2. HgS and Hg2+ bound to sulfur in
              fragments of humic matter.

    The main volatile form in air is elemental mercury but dimethylmercury
    may also occur (Slemr et al., 1951).

         Uncharged complexes, such as HgCl2, CH3HgOH etc., occur in
    the gaseous phase, but are also relatively stable in fresh water (snow
    and rain as well as standing or flowing water). HgCl42- is the
    dominant form in sea water.

    3.3  Levels in the Environment

         The following data have been extracted from Lindquist et al.
    (1984) and are included here to indicate background levels of mercury
    in the environment. Considerable local variations can occur and local
    levels close to anthropogenic sources of mercury would be much higher.

         Reliable data on mercury concentrations in the  air are scarce.
    Recent information suggests a background level at about 2 ng/m3 in
    the lower troposphere of the northern hemisphere and about 1 ng/m3
    in the southern hemisphere, at least over oceanic areas. In European
    areas remote from industrial sources, such as the rural parts of
    southern Sweden and Italy, concentrations most often lie in the range
    from 2 to 3 ng/m3 in summer and from 3 to 4 ng/m3 in winter
    (Brosset 1983, Ferrara et al., 1982). In urban air the concentrations
    could be higher.

         Deposition with  precipitation is a major factor in removing
    mercury from the atmosphere. The lowest concentrations of mercury in
    rain water, around 1 ng/litre, have been reported from a coastal site
    in Japan and from the islands of Samoa. Most other values reported lie
    in the range between 5 and 100 ng/litre.

         Recent measurements of mercury in  aquatic systems have given
    the following concentration ranges, which may be considered
    representative for dissolved mercury:

              Open ocean          0.5-3 ng/litre

              Coastal sea water   2-15 ng/litre

              Rivers and lakes    1-3 ng/litre

         Local variations from these values are considerable, especially
    in coastal sea water and in lakes and rivers where mercury associated
    with suspended material may also contribute to the total load.

         The mercury content in minerals forming ordinary rock and  soils
    is usually very low. The normal level in igneous rocks and minerals
    seems to be less than 50 µg/kg, and in many cases is less than
    10 µg/kg. Due to the strong binding of mercury to soil particles,
    including organic matter, only small amounts of the metal are present
    in soil solution; reported averages range between 20 and 625 µg/kg
    soil.

         Background levels in  sediments are approximately the same as
    levels in unpolluted surface soils. Average concentrations in ocean
    sediments probably lie in the range between 20 and 100 µg/kg.

    3.4  Methylation of Mercury

         The methylation of inorganic mercury in the sediment of lakes,
    rivers and other waterways, as well as in the oceans, is a key step in
    the transport of mercury in aquatic food chains.

         It was first demonstrated by Jensen & Jernelov (1967) that
    microorganisms in lake sediments could methylate mercury. They later
    showed that the degree of methylation correlated well with the overall
    microbial activity in the sediment (Jensen & Jernelov, 1969). Detailed
    mechanisms of methylation in microorganisms have been proposed by Wood
    (1971) and Landner (1971). Some soil organisms capable of methylating
    mercury have also been isolated (Kitamura et al., 1969; Yamada &
    Tonamura, 1972).

         The following general conclusions have been drawn by Bisogni &
    Lawrence (1973) concerning methylation by microorganisms:

         (a)  mono-methylmercury is the predominant product of biological
              methylation near neutral pH,

         (b)  the rate of methylation is greater under oxidising
              conditions than under anaerobic conditions,

         (c)  the output of methylmercury doubles for a ten-fold increase
              in inorganic mercury,

         (d)  temperature affects methylation as a result of its effect on
              overall microbial activity,

         (e)  higher microbial growth rate increases mercury methylation,

         (f)  methylation rates are inhibited by the addition of sulfide
              to anaerobic systems.

         The formation of new or enlarged artificial lakes considerably
    increases the production of methylmercury, although this increase was
    found to be short-lived in new lakes in Finland (Simola & Lodenius,
    1982; Alfthan et al., 1983). A similar problem of increased mercury in
    new lakes, which was taken up by fish and fish-eating mammals,
    occurred in the scheme to divert the Churchill River in Manitoba,
    Canada (Canada-Manitoba, 1987). Methylation rates in one lake, which
    had been flooded 20 years previously, had returned to normal.
    Methylation rates in the new lake, which had flooded arboreal forest,
    were high and were expected to remain high for decades. The source of
    mercury in all of these artificial lakes appeared to be natural rather
    than anthropogenic in origin. Anaerobic conditions after the flooding
    of large amounts of organic material and the subsequent increase in
    microbial activity are thought to be the causes of the increased
    availability of mercury through methylation.

    4.  UPTAKE, LOSS, AND ACCUMULATION IN ORGANISMS

          Background levels of naturally-occuring mercury in the
     environment are generally low, except in the immediate vicinity of
     mining sites and chloralkali plants for the industrial extraction of
     mercury. The majority of mercury in the environment is natural rather
     than the result of human activities. Inorganic mercury can be
     methylated in the environment and the resultant methylmercury is
     taken up into organisms readily; more readily than inorganic mercury.
     Although environmental levels are low, the high capacity of organisms
     to accumulate mercury means that the metal is found widely in both
     aquatic and terrestrial animals and plants. Methylmercury is released
     more slowly by aquatic organisms than inorganic mercury. Aquatic
     invertebrates, and particularly aquatic insects, accumulate mercury
     to a greater extent than fish.

          Speciation of mercury is of great importance in determining the
     uptake of the metal from water and soil. Much of the mercury in
     natural waters and in soil is strongly bound to sediment or organic
     material and is unavailable to organisms.

          Mercury has been found in many terrestrial organisms, birds
     being the subjects of most of the monitoring.

          In many experimental studies, the concentrations of mercury
     quoted are nominal rather than measured. Few attempts have been made
     to estimate available mercury in experimental studies.

          Because of the very extensive literature on the uptake of metals
     into organisms, this section contains illustrative examples and is
     not exhaustive.

         Bioconcentration factors for mercury, determined in laboratory
    experiments, are summarized in Tables 1 and 2.

         Bioconcentration factors are simple ratios between the
    concentration of mercury in an organism and the concentration in the
    medium to which the organism was exposed. This means that results
    should be treated with caution. A relatively low body burden resulting
    from exposure to very low levels of mercury in the medium can give a
    high bioconcentration factor. Conversely, exposure to very high
    mercury levels in the medium can lead to a low bioconcentration
    factor. Exposure to mercury under static test conditions will lead to
    the removal of mercury during the course of the test, whereas flow-
    through conditions maintain a constant level of exposure. Since
    mercury is strongly bound to sediment in the field, it is unclear
    which of these two exposure regimes is the most realistic. It is
    probable that static exposure underestimates and flow-through exposure
    overestimates mercury uptake. Most studies have failed to distinguish


        Table 1.  Accumulation of mercury into aquatic organisms
                                                                                                                                              

    Organism         Lifestagec  Stat/  Organb     Temperature  Compounde                 Duration   Exposure   Bioconcentration    Reference
                                 flowa                (°C)                                 (days)   (µg/litre)      factorg
                                                                                                                                              

    Alga                                                        mercuric chloride           2         164         8537h          Parrish & Carr
    (Croomonas                                                                                                                   (1976)
    salina)

    Filamentous algae            stat                           phenyl mercuric acetate     35        10          1200f          Hannerz (1968)
    (Oedogonium sp.)             flow                  16.5     methoxyethylmercuric OH     18        0.58        2610f          Hannerz (1968)
                                 flow                  15.2     mercuric chloride           54        0.05        871f           Hannerz (1968)

    Duckweed                     flow                           methylmercuric OH           32        3           2950f          Hannerz (1968)
    (Lemna minor)                flow                  16.5     methoxyethylmercuric OH     24        0.58        480f           Hannerz (1968)
                                 flow                  15.2     mercuric chloride           41        0.05        70f            Hannerz (1968)

    Water hyacinth     mature    stat   roots          23-27    mercuric chloride           16        1000        580            Muramoto & Oki
    (Eichhornia                                                                                                                  (1983)
    crassipes)

    Reed                         stat   emergent                phenyl mercuric acetate     35        10          0f             Hannerz (1968)
    (Phragmites                  stat   submerged               phenyl mercuric acetate     35        10          850f           Hannerz (1968)
    communis)                    flow   emergent                methylmercuric OH           32        3           25f            Hannerz (1968)
                                 flow   submerged               methylmercuric OH           32        3           530f           Hannerz (1968)
                                 flow   emergent       16.5     methoxyethylmercuric OH     24        0.58        74f            Hannerz (1968)
                                 flow   submerged      16.5     methoxyethylmercuric OH     24        0.58        139f           Hannerz (1968)
                                 flow   emergent       15.2     mercuric chloride           41        0.05        56f            Hannerz (1968)
                                 flow   submerged      15.2     mercuric chloride           14        0.05        149f           Hannerz (1968)

    Bulrush                      stat   emergent                phenyl mercuric acetate     35        10          90f            Hannerz (1968)
    (Scirpus                     stat   submerged               phenyl mercuric acetate     35        10          790f           Hannerz (1968)
    lucustris)                   flow   emergent                methylmercuric OH           32        3           8f             Hannerz (1968)
                                 flow   submerged               methylmercuric OH           32        3           1250f          Hannerz (1968)
                                                                                                                                              

    Table 1 (cont'd)
                                                                                                                                              

    Organism         Lifestagec  Stat/  Organb     Temperature  Compounde                 Duration   Exposure   Bioconcentration    Reference
                                 flowa                (°C)                                 (days)   (µg/litre)      factorg
                                                                                                                                              

                                 flow   emergent       16.5     methoxyethylmercuric OH     24        0.58        39f            Hannerz (1968)
                                 flow   submerged      16.5     methoxyethylmercuric OH     24        0.58        190f           Hannerz (1968)
                                 flow   emergent       15.2     mercuric chloride           21        0.05        77f            Hannerz (1968)
                                 flow   submerged      15.2     mercuric chloride           41        0.05        70f            Hannerz (1968)

    Yellow iris                  stat   emergent                phenyl mercuric acetate     35        10          20f            Hannerz (1968)
    (Iris                        stat   submerged               phenyl mercuric acetate     35        10          40f            Hannerz (1968)
    pseudacorus)                 flow   emergent                methylmercuric OH           32        3           18f            Hannerz (1968)
                                 flow   submerged               methylmercuric OH           32        3           34f            Hannerz (1968)
                                 flow   emergent       16.5     methoxyethylmercuric OH     18        0.58        31f            Hannerz (1968)
                                 flow   submerged      16.5     methoxyethylmercuric OH     18        0.58        90f            Hannerz (1968)
                                 flow   emergent       15.2     mercuric chloride           49        0.05        18f            Hannerz (1968)
                                 flow   submerged      15.2     mercuric chloride           49        0.05        23f            Hannerz (1968)

    Bloodworm                    stat                           phenyl mercuric acetate     35        10          12 700f        Hannerz (1968)
    (Chironomidae)               flow                           methylmercuric OH           32        3           3070f          Hannerz (1968)
                                 flow                  16.5     methoxyethylmercuric OH     89        0.58        988f           Hannerz (1968)

    Annelid                      stat                           phenyl mercuric acetate     35        10          2030f          Hannerz (1968)
    (Haemopis                    flow                           methylmercuric OH           32        3           450f           Hannerz (1968)
    sanguisuga)                  flow                  16.5     methoxyethylmercuric OH     89        0.58        1148f          Hannerz (1968)

    Annelid                      flow                           methylmercuric OH           32        3           110f           Hannerz (1968)
    (Glossosiphonia              flow                  16.5     methoxyethylmercuric OH     89        0.58        640f           Hannerz (1968)
    complanata)                  flow                  15.2     mercuric chloride           65        0.05        670f           Hannerz (1968)

    Worm                         flow                           methylmercuric OH           32        3           1780f          Hannerz (1968)
    (Oligochaeta)                flow                  16.5     methoxyethylmercuric OH     65        0.58        690f           Hannerz (1968)
                                 flow                  15.2     mercuric chloride           65        0.05        517f           Hannerz (1968)
                                                                                                                                              

    Table 1 (cont'd)
                                                                                                                                              

    Organism         Lifestagec  Stat/  Organb     Temperature  Compounde                 Duration   Exposure   Bioconcentration    Reference
                                 flowa                (°C)                                 (days)   (µg/litre)      factorg
                                                                                                                                              

    Freshwater leech             flow                  15.2     mercuric chloride           65        0.05        534f           Hannerz (1968)
    (Herpobdella
    octoculata)

    Mussel                              WB                      mercuric chloride           4         50          664            Tsuruga (1963)
    (Mytilus edulis)             flow   WB                                                  4         0.06        236f           Hannerz (1968)

    Short-necked clam                   WB                      mercuric chloride           8         50          190            Tsuruga (1963)
    (Venerupis
    philippinarum)

    Pond snail                   stat                           phenyl mercuric acetate     35        10          1280f          Hannerz (1968)
    (Planorbis sp.)              flow                           methylmercuric OH           32        3           3570f          Hannerz (1968)
                                 flow                  16.5     methoxyethylmercuric OH     31        0.58        1970f          Hannerz (1968)
                                 flow                  15.2     mercuric chloride           49        0.05        795f           Hannerz (1968)

    Giant pond snail             stat                           phenyl mercuric acetate     35        10          1800f          Hannerz (1968)
    (Lymnaea                     flow                           methylmercuric OH           32        3           3480f          Hannerz (1968)
    stagnalis)                   flow                  16.5     methoxyethylmercuric OH     31        0.58        1178f          Hannerz (1968)
                                 flow                  15.2     mercuric chloride           14        0.05        297f           Hannerz (1968)

    Snail                        flow                  16.5     methoxyethylmercuric OH     24        0.58        4266f          Hannerz (1968)
    (Physa                       flow                  15.2     mercuric chloride           14        0.05        637f           Hannerz (1968)
    fontinalis)

    Water flea                   stat                           phenyl mercuric acetate     35        10          3570f          Hannerz (1968)
    (Daphnia sp.)

    Cladoceran                   flow                  16.5     methoxyethylmercuric OH     89        0.58        286f           Hannerz (1968)
    (Eurycerus)
                                                                                                                                              

    Table 1 (cont'd)
                                                                                                                                              

    Organism         Lifestagec  Stat/  Organb     Temperature  Compounde                 Duration   Exposure   Bioconcentration    Reference
                                 flowa                (°C)                                 (days)   (µg/litre)      factorg
                                                                                                                                              

    Copepod                      stat   WB             21-26    mercuric chloride           1         0.1         7600           Hirota et al.
    (Acartia clausi)                                                                                                             (1983)
                                 stat   WB             21-26    methyl mercuric chloride    1         0.1         249 000        Hirota et al.
                                                                                                                                 (1983)

    Grass shrimp                        WB                      mercuric chloride           3         1.5         333            Ray & Tripp
    (Palaemonetes                                                                                                                (1976)
    pugio)

    Mayfly             naiad     stat                           phenyl mercuric acetate     35        10          900f           Hannerz (1968)
    (Ephemeridae)      naiad     flow                           methylmercuric OH           32        3           3290f          Hannerz (1968)
                       naiad     flow                  16.5     methoxyethylmercuric OH     24        0.58        680f           Hannerz (1968)
                       larva     flow                  15.2     mercuric chloride           65        0.05        138f           Hannerz (1968)

    Lesser water                 stat                           phenyl mercuric acetate     35        10          4200f          Hannerz (1968)
    boatman                      flow                           methylmercuric OH           32        3           8470f          Hannerz (1968)
    (Corixa sp.)                 flow                  16.5     methoxyethylmercuric OH     89        0.58        740f           Hannerz (1968)
                                 flow                  15.2     mercuric chloride           65        0.05        414f           Hannerz (1968)

    Water boatman                flow                           methylmercuric OH           32        3           2460f          Hannerz (1968)
    (Notonecta                   flow                  16.5     methoxyethylmercuric OH     89        0.58        674f           Hannerz (1968)
    glaaca)                      flow                  15.2     mercuric chloride           65        0.05        483f           Hannerz (1968)

    Midge              larva     flow   WB                      mercuric chloride           30        5.5         19 600         Rossaro et al.
    (Chironomus                                                                                                                  (1986)
    riparius)          pupa      flow   WB                      mercuric chloride           30        5.5         15 600         Rossaro et al.
                                                                                                                                 (1986)
                       adult     flow   WB                      mercuric chloride           30        5.5         7500           Rossaro et al.
                                                                                                                                 (1986)


                                                                                                                                              

    Table 1 (cont'd)
                                                                                                                                              

    Organism         Lifestagec  Stat/  Organb     Temperature  Compounde                 Duration   Exposure   Bioconcentration    Reference
                                 flowa                (°C)                                 (days)   (µg/litre)      factorg
                                                                                                                                              

    Caddisfly          larva     flow                  16.5     methoxyethylmercuric OH     89        0.58        710f           Hannerz (1968)
    (Trichoptera sp.)  larva     flow                  15.2     mercuric chloride           49        0.05        513f           Hannerz (1968)

    Dragonfly          nymph     flow                  16.5     methoxyethylmercuric OH     89        0.58        1296f          Hannerz (1968)
    (Odonata sp.)

    Damselfly          nymph     flow                  16.5     methoxyethylmercuric OH     89        0.58        1186f          Hannerz (1968)
    (Odonata sp.)      nymph     flow                  15.2     mercuric chloride           65        0.05        655f           Hannerz (1968)

    Alderfly           larva     flow                  16.5     methoxyethylmercuric OH     89        0.58        1270f          Hannerz (1968)
    (Sialis lutaria)

    Cranefly           larva     flow                  16.5     methoxyethylmercuric OH     18        0.58        625f           Hannerz (1968)
    (Tipula sp.)       larva     flow                  15.2     mercuric chloride           41        0.05        840f           Hannerz (1968)

    Great diving       imago     flow                  16.5     methoxyethylmercuric OH     89        0.58        800f           Hannerz (1968)
    beetle
    (Dytiscus
    marginalis)

                       larva     flow                  16.5     methoxyethylmercuric OH     89        0.58        3134f          Hannerz (1968)
                       larva     flow                  15.2     mercuric chloride           65        0.05        603f           Hannerz (1968)
                       imago     flow                  15.2     mercuric chloride           65        0.05        862f           Hannerz (1968)

    Pond skater                  flow                  16.5     methoxyethylmercuric OH     89        0.58        754f           Hannerz (1968)
    (Gerris najas)               flow                  15.2     mercuric chloride           65        0.05        431f           Hannerz (1968)

    Aquatic saw bug              flow                  16.5     methoxyethylmercuric OH     89        0.58        954f           Hannerz (1968)
    (Asellus
    aquaticus)
                                                                                                                                              

    Table 1 (cont'd)
                                                                                                                                              

    Organism         Lifestagec  Stat/  Organb     Temperature  Compounde                 Duration   Exposure   Bioconcentration    Reference
                                 flowa                (°C)                                 (days)   (µg/litre)      factorg
                                                                                                                                              

    Water spiders                flow                  16.5     methoxyethylmercuric OH     89        0.58        624f           Hannerz (1968)
    (Hydracnidae)

    Pike                         stat   liver          17.2     methoxyethylmercuric OH     10        0.4         7673f          Hannerz (1968)
    (Esox lucius)                stat   kidney         17.2     methoxyethylmercuric OH     10        0.4         7230f          Hannerz (1968)
                                 stat   liver                   methylmercuric OH           10        0.3         2002f          Hannerz (1968)
                                 stat   kidney                  methylmercuric OH           10        0.3         2198f          Hannerz (1968)

    Rainbow trout      juv       flow   WB             5        methyl mercuric chloride    84        0.263       4525           Reinert et al.
                                                                                                                                 (1974)
    (Salmo gairdneri)  juv       flow   WB             10       methyl mercuric chloride    84        0.258       6628           Reinert et al.
                                                                                                                                 (1974)
                       juv       flow   WB             15       methyl mercuric chloride    84        0.244       8033           Reinert et al.
                                                                                                                                 (1974)
                                        WBd            5        mercuric chloride           4         50          5              MacLeod &
                                                                                                                                 Pessah (1973)
                                        WBd            10       mercuric chloride           4         50          12             MacLeod &
                                                                                                                                 Pessah (1973)
                                        WBd            20       mercuric chloride           4         50          26             MacLeod &
                                                                                                                                 Pessah (1973)

    Bluegill sunfish             stat   WB             9        methyl mercuric chloride    28.6      0.5         222f           Cember et al.
                                                                                                                                 (1978)
                                                                                                                                              

    Table 1 (cont'd)
                                                                                                                                              

    Organism         Lifestagec  Stat/  Organb     Temperature  Compounde                 Duration   Exposure   Bioconcentration    Reference
                                 flowa                (°C)                                 (days)   (µg/litre)      factorg
                                                                                                                                              

    (Lepomis                     stat   WB             21       methyl mercuric chloride    28.6      0.5         1138f          Cember et al.
    macrochirus)                                                                                                                 (1978)
                                 stat   WB             33       methyl mercuric chloride    28.6      0.5         2454f          Cember et al.
                                                                                                                                 (1978)
                                                                                                                                              

    a  stat = static conditions (water unchanged for duration of experiment); flow = flow-through conditions (mercury concentration in water
       continuously maintained).
    b  WB = whole body.
    c  juv = juvenile.
    d  muscle, skin & bone.
    e  OH = hydroxide.
    f  radiometrically calculated.
    g  bioconcentration factor = concentration in organism/concentration in medium.
    h  dry weight.

    Table 2.  Accumulation of mercury into terrestrial organisms
                                                                                                                                           

    Organism              Route   Organ     Compound                         Duration   Exposureb   Bioconcentration   Reference
                                                                             (days)     (mg/kg)        factorc
                                                                                                                                           

    Broccoli              soil    leaves    mercuric chloride                60         20              0.002d        John (1972)
    (Brassica oleracea)   soil    roots     mercuric chloride                60         20              0.09d         John (1972)

    Pea                   soil    roots     mercuric chloride                95         20              0.07d         John (1972)
    (Pisum sativum)

    Cauliflower           soil    leaves    mercuric chloride                70         20              0.003d        John (1972)
    (Brassica oleracea)   soil    roots     mercuric chloride                70         20              0.12d         John (1972)

    Spinach               soil    leaves    mercuric chloride                55         20              0.03d         John (1972)
    (Spinacia oleracea)   soil    roots     mercuric chloride                55         20              0.05d         John (1972)

    Chicken               diet    muscle    methyl mercury dicyandiamide     35-42      8               1.25           Borg et al. (1970)
                          diet    liver     methyl mercury dicyandiamide     35-42      8               5              Borg et al. (1970)

    Mallard               diet    liver     methyl mercury dicyandiamide     14         8               2.1            Stickel et al. (1977)
    (Anas platyrhynchos)  diet    kidney    methyl mercury dicyandiamide     14         8               2.2            Stickel et al. (1977)

    Redwinged blackbird   diet    liver     methyl mercury dicyandiamide     11         40              2.3            Finley et al. (1979)
    (Agelaius             diet    kidney    methyl mercury dicyandiamide     11         40              2.1            Finley et al. (1979)
    phoeniceus

    Cowbird               diet    liver     methyl mercury dicyandiamide     11         40              1.7            Finley et al. (1979)
    (Molothrus ater)      diet    kidney    methyl mercury dicyandiamide     11         40              1.5            Finley et al. (1979)

    Grackle               diet    liver     methyl mercury dicyandiamide     11         40              1.3            Finley et al. (1979)
    (Quiscalus quiscula)  diet    kidney    methyl mercury dicyandiamide     11         40              1.1            Finley et al. (1979)
                                                                                                                                           

    Table 2 (cont'd)
                                                                                                                                           

    Organism              Route   Organ     Compound                         Duration   Exposureb   Bioconcentration   Reference
                                                                             (days)     (mg/kg)        factorc
                                                                                                                                           

    Mink                  diet    liver     ceresan La                       5          32              11.1           Aulerich et al.(1974)
    (Mustela vision)      diet    kidney    ceresan La                       5          32              7.4            Aulerich et al.(1974)
    (adult)               diet    liver     mercuric chloride                10         135             0.3            Aulerich et al.(1974)
                          diet    kidney    mercuric chloride                10         135             3.2            Aulerich et al.(1974)
                                                                                                                                           

    a  ceresan L = methylmercury 2,3-di-hydroxy propyl mercaptide + methylmercury acetate.
    b  exposure as mg/kg of mercury soil or diet according to route.
    c  concentration factors calculated on a wet weight basis unless otherwise stated; bioconcentration factor = concentration in
       organism/concentration in medium.
    d  dry weight.
    
    between mercury taken into the tissues of the organism and mercury
    adsorbed on external surfaces. This should also be taken into account
    when interpreting results.

         Taking these factors into account, it is still clear that
    organisms take up both inorganic and organic forms of mercury from the
    medium. This uptake can result in high concentration factors. Under
    identical conditions, organic mercury is taken up by organisms to a
    greater degree than inorganic mercury, although the latter may often
    be strongly adsorbed to the outer surfaces.

    4.1  Speciation of Mercury

    Appraisal

          Different species of mercury differ greatly in their
     physicochemical properties: in their solubility, rates of
     accumulation by organisms, and behaviour in ecosystems. It is in its
     methyl form that mercury is most hazardous. Although not all sites of
     methylation in the environment are fully known, several have been
     identified in the aquatic environment.

         Mercury accumulated in the tissues of fish is usually in the form
    of methylmercury, while the source is usually inorganic mercury
    (Huckabee et al., 1979). Several hypotheses of how and where
    methylation occurs have been proposed. The main hypotheses are:

         (a)  biological methylation, bacterial in origin, which produces
              methylmercury in the environment (methylmercury is taken up
              by fish more readily than inorganic mercury),

         (b)  methylation by microorganisms associated with branchial
              mucus of the fish or in the fish gut, and

         (c)  methylation in the fish's liver (Thellen et al, 1981).

    It is generally agreed that methylation by fish, other than by
    bacteria associated with the fish, either does not occur or accounts
    for only an insignificant amount of the methylmercury produced. There
    is good evidence for methylation by bacteria in aquatic systems.

         Jernelov (1968) suggested that fish could not methylate mercury
    themselves and this is generally accepted (Huckabee et al., 1979),
    though not universally. Jernelov & Lann (1971) showed that 60% of the
    mercury content of predator fish (northern pike) arose from prey fish.
    This mercury was already methylated in the prey. The concentration of
    mercury in predator species was similar to that in their prey. They
    also measured the mercury content of organisms that were the food of
    the prey fish. Mercury levels in benthic fauna were very low and
    contributed less than 25% of the mercury in bottom-feeding fish. Most
    of the mercury accumulated by non-predator species was, therefore,

    accumulated directly from water. This conclusion was also reached by
    Fagerstrom & Asell (1973). The question of where the methylation,
    which gives rise to methylmercury residues in fish, occurs is still
    unresolved. It is also generally accepted that fish do not demethylate
    mercury either.

    4.2  Uptake and Loss in Aquatic Organisms

    Appraisal

          The data presented on uptake by aquatic invertebrates are
     difficult to interpret because most studies do not differentiate
     between external adsorption and actual uptake into the organism. This
     is especially important for methylmercury compounds for which uptake
     seems to be correlated with surface adsorption capacity, as expressed
     by the relative size of the organism.

          The extrapolation of data on uptake to other organisms appears
     risky because of a lack of knowledge regarding the mechanisms of
     uptake. This is even true for phenomena that are apparently fairly
     universal, e.g., the facilitating influence of chelators upon
     uptake.

          Most data on uptake by fish support the notion that uptake
     correlates positively with available concentration, with exposure
     time, and with temperature, although hardly any investigation
     differentiates between nominal and available concentrations. The
     importance of this distinction seems to be illustrated by the
     positive influence of lowered pH upon uptake.

          None of the studies address the problem of distinguishing
     between adsorption to gills and slime on the one hand and real uptake
     into the body on the other. Studies of mercury distribution between
     organs are valuable for the potential effects of the total body
     burden, but they give no reliable insight into the time-dependent
     process of accumulation.

          Data consistently show a higher uptake of methylmercury than of
     inorganic mercury. However, other organic mercury compounds exhibit a
     lower uptake, since they are adsorbed to a lesser extent.

    4.2.1  Microorganisms, plants, and invertebrates

         When Glooschenko (1969) exposed the marine diatom  Chaetoceros
     costatum to labelled mercury, he found no difference between uptake
    in the light or the dark in non-dividing cells. Dead cells took up
    twice as much mercury as living cells, presumably by surface
    adsorption. As dividing cells in the light accumulated the labelled
    mercury for longer than non-dividing cells, the author suggested the
    possibility of some active uptake.

         Hannerz (1968) demonstrated that there was no appreciable
    assimilation of mercury into the tissues of aquatic plants. Although
    concentrations were 10-20 times higher in submerged parts compared to
    emergent parts, this was attributed to surface adsorption differences.
    De et al. (1985) grew the plant  Pistia stratiotes in nutrient
    solution to which mercuric chloride had been added at concentrations
    ranging from 0.05 to 20 mg/litre. They found that uptake gradually
    increased with an increase in the mercury concentration. Maximum
    accumulation occurred within one day. Maximum removal (approximately
    90%) was recorded at 6 mg/litre or less, only 20% being lost from
    plants receiving the highest concentration. Mercury accumulation into
    the roots was about 4 times higher than into the shoots at lower
    concentrations and about twice as high at 20 mg/litre.

         Zuberik & O'Connor (1977) studied the accumulation of mercury in
    aquatic organisms from the Hudson River, USA. The organisms were
    maintained in filtered river water that contained mercury
    concentrations of < 0.1 µg/litre (less than levels normally found in
    the Hudson River). Planktonic organisms were exposed to various forms
    of labelled mercury, and the concentration factors after 24 h ranged
    from 102 to 106. Mercury uptake was greater in microplankton and
    algae than in macroplankton and fish larvae. An amphipod  (Gammarus
     sp.) was exposed for one day to each of four types of mercury, two
    organic (phenylmercuric acetate and methylmercury chloride) and two
    inorganic (mercurous nitrate and mercuric chloride). No differences in
    uptake were found, but when the amphipod was exposed for a week the
    organic forms were accumulated to 3 times the concentration of the
    inorganic forms.

         Riisgard et al. (1985) transferred mussels  (Mytilus edulis)
    from clean water to an area chronically polluted with mercury. The
    mussels accumulated mercury readily during 3 months of exposure. They
    were then transferred to clean water in the laboratory and the
    elimination of the mercury was measured. The biological half-life was
    293 days, but was only 53 days in the case of mussels contaminated by
    a temporary massive mercury contamination. In both cases, 75% of the
    mercury in the mussels was inorganic, but both inorganic and organic
    species were immobilized in the mussels from the chronically polluted
    area. In another study, only 6% of the total mercury in  Macoma
     balthica, a sediment-feeding bivalve, was methylated, a much lower
    percentage than in  Mytilus from the same area.

         Hirota et al. (1983) exposed the copepod  Acartia clausi to
    inorganic (mercuric chloride) and organic (methylmercury chloride)
    mercury at concentrations of 0.05-0.5 µg/litre for 24 h. The
    bioconcentration factor for inorganic mercury was nearly constant
    (approximately 7500), regardless of the mercury concentration in the
    water or the density of the copepods. In contrast, the concentration
    factor of methylmercury fluctuated, showing an inverse relationship
    with density but no relationship with the mercury concentration in the
    water.

         DeFreitas et al. (1981) found a net assimilation of 70%-80% for
    methylmercury and 38% for inorganic mercury when fed in the diet to
    the shrimp  Hyalella azteca. From water, inorganic mercury was
    assimilated 2 to 3 times more slowly than methylmercury. Khayrallah
    (1985) found that the accumulation of ethylmercuric chloride was
    almost twice as rapid as that of mercuric chloride in the amphipod
     Bathyporeia pilosa, although death occurred at similar levels of
    mercury.

         Ray & Tripp (1976) exposed the grass shrimp  (Palaemonetes pugio)
    to radioactively labelled methylmercury chloride and mercuric chloride
    for 24 and 72 h. After 24 h, the methylated form was mostly
    concentrated in the ventral nerve cord and to a lesser extent in the
    gills. The reverse was true for mercuric chloride. The concentrations
    of mercury accumulated in the other tissues (exoskeleton, foregut, and
    remainder) were similar for both compounds, and were in decreasing
    order of the above list. After 72 h the tissue distribution had
    changed, and there was no consistent order of the relative tissue
    concentrations. There was an increase in the mercury levels of the
    exoskeleton, foregut, and remainder tissues, while that in the gills
    remained about the same and that in the ventral nerve cord decreased.

         Vernberg & O'Hara (1972) measured the uptake of labelled mercury
    into the gills and hepatopancreas of fiddler crabs  (Uca pugilator)
    maintained in a solution containing 0.18 mg mercury/litre (as mercuric
    chloride) for 72 h. Uptake was determined under various temperature
    (5°C to 33°C) and salinity (5 and 30 g/litre) regimes. The total
    mercury taken up by the gills and hepatopancreas pooled together was
    unaffected by the different regimes. However, the ratio of uptake into
    the two tissues was affected. At higher temperatures, the crabs seem
    able to transport mercury from gill tissue to the hepatopancreas more
    effectively than at low temperatures.

         When Rossaro et al. (1986) exposed various life stages of the
    midge  Chironomus riparius to mercuric chloride for a period of
    30 days, the levels were still increasing at the end of the
    experiment. Both larvae and pupae accumulated mercury to about the
    same levels, some accumulation being due to passive adsorption. In a
    small experiment to illustrate this, larvae kept in a solution of
    5 µg/litre for only 1 min accumulated 9.32 mg mercury/kg. The adults
    accumulated only 40% of the levels found in the larval stage. The
    authors suggested that this is because the adults have some means for
    eliminating the mercury.

         Getsova & Volkova (1964) reported concentration factors for the
    accumulation of radioactively labelled mercury in four insect species.
    A midge,  Glyphotaelius punctatolineatus, accumulated 5240 times the
    water concentration within 16 days, while a dragonfly,  Leucorrhinia
     rubicunda, accumulated 8310 times the concentration over 16 days.
    Another dragonfly,  Aeschna grandis, accumulated 4000 times the
    waterborne mercury in 8 days, while a waste-water inhabiting fly,

     Eristalis tenax, accumulated only 640 times the water concentration
    after 4 days and the concentration factor had fallen to just 266 after
    8 days. The authors stated that the concentration factors that they
    found were in agreement with other Russian work on mercury
    accumulation.

    4.2.2  Fish

         When Birge et al. (1979) exposed rainbow trout eggs to an
    inorganic mercury concentration of 0.1 µg/litre in a flow-through
    system, the eggs accumulated 42.4, 68.2, and 96.8 µg mercury/kg after
    1, 4, and 7.5 days, respectively. Control eggs contained
    18.6 µg mercury/kg. The bioconcentration factor over 7.5 days was 782,
    taking into account the degree of contamination of controls. This
    represented a daily uptake rate of about 20 µg/kg. There was no
    evidence to suggest that the mercury penetrated the outer covering of
    the eggs and there was a high probability that most of the "uptake"
    was surface adsorption.

         Backstrom (1969) found that the uptake by fish of various mercury
    compounds was similar to that observed with birds (where methylmercury
    is rapidly absorbed compared with phenylmercury, methoxyethylmercury,
    and inorganic mercury), but the difference in uptake between
    methylmercury and the other mercury compounds was less pronounced.
    Mercury uptake into the spleen and the thyroids was greater than for
    birds. Phenylmercury was also retained in the wall of the gall
    bladder. In general the uptake of mercury into fish was far more
    localized than in birds. The levels of methylmercury steadily
    increased in the muscles and in the brain, whereas the other compounds
    accumulated primarily in the kidneys, spleen, and liver. More mercury
    accumulated in red flesh than white. There was also a high uptake of
    mercury into the gills and pseudobranch.

         Kramer & Neidhart (1975) demonstrated that methylmercury was
    taken up from water by guppies  (Lebistes reticulatus) 17 times
    faster than inorganic mercury. Organic mercury was also eliminated
    more slowly than inorganic. The authors suggested that some
    methylation of mercury occurred in the fish.

         Ribeyre & Boudou (1984) examined the uptake of mercury over time
    into specific organs of the rainbow trout. The uptake was sigmoid with
    a linear phase and a plateau. The majority (55% for inorganic and 60%
    for methylmercury) of the metal was found in muscle and gills, while
    blood contained 3%-12%, liver 2%-5%, and kidneys 2%-7%. Brain,
    posterior intestine, and spleen together accounted for only 2% of
    total mercury. Those organs which would eventually contain most
    mercury accumulated their mercury exponentially. After the exposure,
    some organs lost their mercury while others (the ones with most
    mercury) continued to increase their mercury content. The organs which
    lost mercury in clean water had accumulated the metal with a flatter
    sigmoid curve.

         Schindler & Alberts (1977) found that the mosquitofish  (Gambusia
     affinis) readily accumulated metallic mercury during short-term
    continuous exposure. Within 24 h, 20 mg/kg wet weight had been taken
    up from a solution containing 0.1 mg total mercury/litre. The uptake
    curves for metallic mercury and mercuric chloride were very similar.
    The authors suggested that uptake in the short-term is largely the
    result of physical adsorption. This rate of uptake closely agrees with
    that found by McKone et al. (1971) in goldfish  (Carassius auratus)
    where 22 mg mercuric chloride/kg was accumulated from a solution
    containing 0.25 mg/litre over a period of 24 h.

         When Schindler & Alberts (1977) periodically exposed (2 h/day for
    10 days) mosquitofish to metallic mercury and mercuric chloride
    (in separate experiments) at 100 µg/litre, the uptake of metallic
    mercury was 5 times greater than that of the chloride. The authors
    suggested that the metallic mercury remained unchanged and that its
    high lipid solubility enabled it to penetrate the gill membrane,
    whereas the salt bound more tightly to the mucoproteins of the gills
    and penetration was restricted. The rate of elimination in mercury-
    free water was about the same for both, with the half-time calculated
    to be about 45 days.

         McKim et al. (1976) exposed 3 generations of brook trout
     (Salvelinus fontinalis) to methylmercury at concentrations measured
    at < 0.01-2.93 µg/litre. The uptake was rapid and 2-week
    concentration factors ranged from 1000 to 12 000, depending on the
    tissue. There was a tendency for the uptake to reach a steady state
    (that is the tissue content reached a constant level) over 20-28
    weeks. There was no significant elimination over this period.

         In studies by Pentreath (1976), the thornback ray  (Raja clavata)
    readily absorbed both inorganic mercuric chloride and organic
    methylmercuric chloride from sea water. Methylmercury, in contrast to
    inorganic mercury, was readily absorbed from food and slowly
    eliminated. The half-lives of elimination of mercury taken up from
    food were 61.6 days for inorganic and 323 days for organic components.

         Thellen et al. (1981) found that methylmercuric chloride rapidly
    accumulated in the organs and muscular tissue of rainbow trout exposed
    to 1 mg/kg diet. However, mercuric chloride, at the same
    concentration, did not accumulate. During exposure to a continuous
    sublethal concentration of 0.25 µg mercury/litre, both organic and
    inorganic mercury accumulated, primarily in the internal organs and to
    a lesser extent the muscle tissue. Mercuric chloride was detected in
    the muscle at half of the concentration of organic mercury. Wobeser
    (1975b) fed rainbow trout fingerlings a diet containing methylmercuric
    chloride (at 4, 8, 16, or 24 mg mercury/kg) over a 15-week period. The
    total accumulation of mercury in muscle tissue was directly related to
    the concentration of mercury in the food, as was the rate of
    accumulation. Mercury was accumulated in muscle to a higher
    concentration than there had been in the diet.

         When Amend (1970) exposed juvenile sockeye salmon (1 h per day
    for 12 to 15 days) to 1 mg/litre of lignasan (6.25% ethylmercury
    phosphate), the fish contained highest levels in the kidneys and
    liver. One week after the cessation of treatment, these levels were
    36.5 and 20.4 mg/kg for the kidney and liver, respectively. Three
    years later, the fish having migrated, levels were still higher than
    normal but had returned to normal after 4 years. Similar studies using
    coho and chinook salmon yielded similar results. When Kendall (1975)
    injected channel catfish intraperitoneally with methylmercury chloride
    at 15 mg/litre, the mean concentration of mercury in the kidneys was
    51.03 µg/g after 24 h and fell to 14.24 mg/kg after 96 h.

    4.2.2.1  Effects of environmental variables on uptake by fish

    Appraisal

          Environmental variables such as temperature and pH increase the
     uptake of mercury, particularly methylmercury, by fish. This is of
     potentially considerable importance in the field.

         Reinert et al. (1974) found that yearling rainbow trout  (Salmo
     gairdneri) exposed to methylmercury chloride for 12 weeks accumulate
    more mercury at 15°C than at 5°C (Table 1). When Cember et al. (1978)
    exposed bluegill sunfish  (Lepomis macrochirus) to methylmercury
    chloride at concentrations ranging from 0.2 to 50 µg/litre for up to
    688 h, mercury accumulation was not affected by the different mercury
    concentrations. It did, however, increase when the temperature was
    increased from 9°C to 33°C (Table 1). MacLeod & Pessah (1973) found an
    increase in mercury accumulation, in response to an increase in
    temperature (from 5 to 20°C), in rainbow trout exposed to
    concentrations of between 50 and 200 µg/litre for 4 days. The authors
    also interpolated (from 7-day data) a 4-day bioconcentration factor
    for phenylmercuric acetate of 100, when the fish were exposed to
    5 µg/litre mercury at 10°C. Tsai et al. (1975) studied the effect of
    pH on the accumulation of inorganic mercury (mercuric chloride) at a
    concentration in water of 1500 µg mercury/litre for 15 min. The
    accumulation increased as pH decreased. At pHs of 5, 6.5, and 7.5,
    fathead minnow accumulated whole body residues of 2.7, 1.8, and
    0.4 mg mercury/kg, calculated on a wet weight basis, respectively. A
    similar result was found for the emerald shiner  (Nicropterus
     atherinoides).

         Rodgers & Beamish (1981) found that the uptake of methylmercury
    by rainbow trout was increased when the hardness of the water was
    decreased from 385 mg/litre to 30 mg/litre. The addition of inorganic
    mercuric chloride increased the uptake of methylmercury in both hard
    and soft water. Kudo & Mortimer (1979) exposed guppies to mercury in a
    double chambered system, with an exchange of water. Only in one
    chamber did the fish have access to sediment. After being exposed for

    20 days to a sediment mercury concentration of 1.023 mg/kg, the fish
    without direct access to the sediment showed a concentration factor of
    57 and those with access a factor of 570.

    4.2.3  Studies on more than one type of organism

         Cultures of the alga  Croomonas salina, grown for 48 h in the
    presence of mercuric chloride (164 µg mercury/litre), retained about
    half of the mercury (1400 mg/kg dry weight) (Parrish & Carr 1976).
    When the alga was fed to the copepod  Acartia tonsa for 5 days,
    neither the copepods nor their eggs or faeces retained mercury in
    detectable amounts.

         Boudou et al. (1979) exposed mosquitofish  (Gambusia affinis) to
    methylmercury directly from the water and via food organisms and water
    in a simple model ecosystem. More mercury was taken up at higher
    temperatures. The authors calculated mercury uptake from water as a
    percentage of the "global" uptake from both water and food. This
    percentage varied with temperature, being 83% at 10°C, 40% at 18°C,
    and 11% at 26°C.

         In studies by Boudou & Ribeyre (1984), alevins of rainbow trout
     (Salmo gairdneri) were exposed to a constant water concentration of
    methylmercuric or mercuric chloride at 1 µg/litre for 83 days. Mercury
    uptake was faster with organic than inorganic and both were initially
    linear. A plateau was eventually achieved in both cases. Uptake was
    negatively related to fish weight, although the authors pointed out
    that in the field there is usually a positive relationship.

         Fang (1973) maintained the pond weed  Elodea canadensis, snail
     Helisoma campanulata, coontail plant  Ceratophyllum demersum, and
    guppy  Lebistes reticulatus in solutions containing labelled
    phenylmercuric acetate (PMA) at concentrations between 5 × 10-8 and
    5 × 10-7mol/litre. All of the organisms readily accumulated PMA and
    the uptake was related to the length of exposure and the
    concentration. The absorbed PMA was largely converted to inorganic
    mercury. Although the uptake curves were very similar, pond weed and
    coontail both accumulated much more PMA than guppy or snail. The half-
    life of Hg203 residues ranged from 43 to 58 days. When Fang (1974)
    exposed  Lebistes reticulatus and  Ceratophyllum demersum to
    labelled ethylmercuric chloride (EMC), the uptake was positively
    related to the time of exposure over 200 h and the concentration up to
    5 × 10-7mol/litre. Highest concentrations were accumulated in the
    internal organs. The half-life of EMC was 20-23 days. Both organisms
    converted EMC to inorganic mercury, 34% being converted by the
    coontail and 29% by the guppy over a 7-day period. When the same
    organisms were exposed to methylmercury chloride, little or no
    breakdown to inorganic mercury occurred.

    4.3  Uptake and Loss in Terrestrial Organisms

    Appraisal

          The accumulation of mercury in plants increases with increasing
     soil mercury concentration. Soil type has a considerable influence on
     this process, a high organic matter content decreasing the uptake.
     Generally, the highest concentrations of mercury are found at the
     roots, but translocation to other organs (e.g., leaves) occurs. In
     contrast to higher plants, mosses take up mercury via the
     atmosphere.

          In exposed birds, the highest mercury levels are generally found
     in liver and kidneys. Methylmercury is more readily absorbed than
     inorganic mercury and it exhibits a longer biological half-time.
     Depending on speciation, mercury occurs in different compartments of
     birds' eggs; methylmercury tends to concentrate in the white and
     inorganic mercury in the yolk.

         Huckabee & Janzen (1975) found that the mat-forming moss
     Dicramum scoparium did not take up radioactively labelled mercury
    from substrate. The authors concluded that the uptake of mercury into
    this point was mostly from the atmosphere. This is commonly true for
    mosses, which have been used extensively as monitor organisms for
    atmospheric pollutants in the field. Weaver et al. (1984) maintained
    bermuda grass  (Cynodon dactylon) in three types of soil (clay, silt
    loam, and fine sand) treated with mercuric chloride (1-50 mg/kg).
    Mercury was accumulated into the roots from silt loam, clay, and sand
    in increasing order. The accumulation increased with increasing
    mercury concentration. At 50 mg/kg the concentration of mercury in
    (and on the surface of) the roots was 800 mg/kg, when the grass was
    grown in sand.

         John (1972) grew eight types of food crop in soil treated with
    mercuric chloride at 4 or 20 mg mercury/kg, and uptake was measured
    after 35 to 130 days, depending on the plant species. Higher
    concentrations of mercury were found in the roots compared to the
    above-ground samples. At the highest treatment level the mercury
    content of the roots, calculated on a dry weight basis, ranged from
    0.387 mg/kg for lettuce to 2.447 mg/kg for cauliflower. Of the edible
    plant parts, spinach leaves and radish tubers contained the highest
    concentrations (0.695 and 0.663 mg/kg mercury, respectively).

         Siegel & Siegel (1985) found that the seed-pods of several
    leguminous species exposed to soil mercury concentrations of
    10-69 µg/kg lost 75-85% of their tissue water during maturation but
    showed no loss of mercury content. However, the seeds not only lost
    most of their water but also at least 75% of their mercury. The
    authors suggested that the elimination was by "bio-volatilisation",
    i.e., loss of elemental mercury as vapour rather than by
    translocation.

         Nuorteva et al. (1980) reared blowfly  (Lucilia illustris) on
    trout flesh contaminated with mercury (0.66 mg/kg). Levels rose from
    0.14 to 1.18 mg/kg during the larval feeding period, whereas pupae and
    freshly emerged adults contained 0.99 and 1.01 mg/kg, respectively.
    When adults were then fed honey, mercury levels were reduced to a
    third within 2 days. The authors found that it was easier for the
    flies to eliminate inorganic mercury than methylmercury. Nuorteva &
    Nuorteva (1982), after rearing blowfly larvae on mercury-contaminated
    fish flesh and obtaining mercury levels of 2, 6.3, and 13.3 mg/kg in
    different groups, fed the flies to staphylinid beetles  (Creophilus
     maxillosus) for a 1-week period. This gave residues of 6.9, 17.4,
    and 33.4 mg/kg, respectively, in the beetles.

         Kiwimae et al. (1969) fed white leghorn hens for 140 days on a
    diet containing 400 or 1600 µg of mercury per day as either mercury
    nitrate, phenylmercury hydroxide, or methoxyethylmercury hydroxide.
    The total mercury accumulated in the egg-whites of eggs laid was 0.31,
    0.53, and 0.46 mg/kg, respectively, for the lower dose and 0.44, 0.85,
    and 0.88 mg/kg for the higher dose. At the higher dose, the mercury
    residue in the egg yolks was 2.12, 4.53, and 2.89 mg/kg, for the three
    mercury compounds, respectively.

         Backstrom (1969) administered labelled mercury compounds, either
    parenterally or perorally, to Japanese quail and studied the tissue
    uptake and elimination. The route of administration did not affect the
    final uptake or subsequent elimination. Methylmercury was readily
    absorbed and was stable, while the other compounds, phenylmercury,
    methoxyethylmercury, and inorganic mercury, were less well absorbed,
    and the phenylmercury was rapidly decomposed to inorganic mercury.
    Methylmercury was characterized by an even tissue distribution and a
    slow excretion, which was enhanced in egg-laying hens. The author
    attributed this to an increased concentration of methylmercury in the
    egg-white. Little of the other compounds were taken up into the brain,
    but methylmercury slowly reached a high concentration. The other
    mercury compounds were accumulated in the  yolks of the eggs laid,
    and also in the liver and kidneys of the adult birds, and were rapidly
    excreted. The plumage and other keratinised structures strongly
    concentrated mercury, irrespective of the compound. These structures
    seem to be an important excretion route, especially for methylmercury.

         Nicholson & Osborn (1984) fed juvenile starlings  (Sturnus
     vulgaris) on a mercury-contaminated synthetic diet
    (1.1 mg mercury/kg) and analysed the birds after 8 weeks. The highest
    mercury levels were found in the kidneys and the liver (36.3 and
    6.55 mg/kg dry weight, respectively).

         In studies by Finley & Stendell (1978), black ducks  (Anas
     rubripes) were fed a diet containing 3 mg mercury/kg (as
    methylmercury dicyandiamide) for periods of 28 weeks over two
    consecutive breeding seasons, during which time any ducklings that
    hatched were also fed the dosed diet. Mercury levels were highest in

    the feathers of the adult birds (61 mg/kg wet weight), followed by the
    liver and kidneys (22 and 14 mg/kg, respectively). Similarly the
    highest levels were also found in the feathers, liver, and kidneys of
    first-year ducklings. Eggs and embryos analysed during the first year
    revealed mercury levels of 6.14 and 9.62 mg/kg, respectively. Mercury
    residues in eggs, embryos, and ducklings were, on average, about 30%
    lower during the second year. Stickel et al. (1977) dosed mallard
     (Anas platyrhynchos) with 8 mg mercury/kg for 2 weeks, and found
    that the highest levels of mercury were accumulated in the liver
    (16.5 mg/kg wet weight) and the kidney (17.6 mg/kg wet weight). One
    week later the liver and kidney had retained 64 and 66%, of the
    mercury, respectively. No significant additional loss was noted during
    the next 8 weeks.

         Adams & Prince (1976) showed that ring-necked pheasants
     (Phasianus colchicus) accumulated more mercury in the tissues after
    consuming methylmercury dicyandiamide than after consuming the
    corresponding mass of phenylmercuric acetate. This reflects the
    greater toxicity of alkyl mercury compounds than aryl ones.

         When Borg et al. (1970) fed goshawks  (Accipiter gentilis) liver
    and muscle from chickens dosed with methylmercury (average dietary
    mercury content 13 mg/kg), the hawks died within 6-7 weeks. The
    highest residues of mercury were found in the liver at 113 mg/kg wet
    weight (102 mg methylmercury/kg), and the kidneys at 129 mg/kg
    (98 mg methylmercury/kg). Substantially higher levels of mercury were
    found in the skeletal muscle and brain of treated birds than in those
    of controls. The reproductive organs also showed an ability to
    accumulate mercury.

    4.4  Accumulation in the Field

    Appraisal

          Observations on given species of marine and freshwater fish
     indicate that all tissue concentrations of mercury increase with
     increasing age (as inferred from length) of the fish. In certain
     species males have been found to have higher levels than females.

          In aquatic systems, fish-eating birds tend to have higher
     mercury levels than non-fishing birds. In terrestrial systems, seed-
     eating birds, small mammals, and their predators can have high levels
     in areas where methylmercury fungicides are used.

          Bird feathers are useful for biological monitoring for
     methylmercury exposure. Analysis of feathers, especially using
     neutron activation, can allow recapitulation of past exposure. In
     general liver and kidney have higher levels than other bird tissues.

          Sea mammals are reported to have a wide range of total mercury
     concentrations in liver (0.4 to over 300 mg/kg), only a small
     fraction (2-17%) being in the methylated form. Selenium and mercury
     have been found in seal livers in a consistent 1:1 atomic ratio. A
     number of studies have indicated that selenium plays a protecting
     role.

          Point sources of mercury pollution often lead to elevated
     mercury levels in organisms living in the affected area. There are
     some circumstances where toxic effects have been produced. These
     effects should be taken into account in various countries during the
     process of industrialization.

    4.4.1  General exposure

         Gilmartin & Revelante (1975) analysed Northern Adriatic anchovy
     (Engraulis encrasicholus) and sardine  (Sardina pilchardus) for
    mercury content. Seasonal distribution of mercury in various tissues
    of both anchovy and sardine ranged between 5 and 610 ng/g wet weight,
    the highest concentrations of mercury being in the liver and kidney.
    Perttila et al. (1982) found that mercury levels in the Baltic herring
     (Clupea harengus) increased significantly with age. Bache et al.
    (1971) observed that concentrations of both total mercury and
    methylmercury increased with the age of lake trout  (Salvelinus
     namaycush), the proportion of methylmercury to total mercury
    increasing with age. However, Westoo (1973) did not find that the
    proportion of methylmercury to total mercury in salmon  (Salmo salar)
    and sea trout  (Salmo ocla) was dependant on age.

         Forrester et al. (1972) found a correlation between length and
    mercury concentration in  Squalus acanthias (the spurdog, an
    elasmobranch fish). Olsson (1976) analysed northern pike  (Esox
     lucius) in 1968 and 1972 and found a correlation between mercury
    levels and length of fish, and that males contained significantly more
    mercury than females. It was considered that, during a general
    decrease of mercury levels within pike population, the age of the fish
    is not a suitable parameter for estimating mercury levels. This is
    because uptake and retention of mercury is dependant on body size but
    loss of accumulated mercury is less dependent on fish size. May &
    Mckinney (1981) sampled freshwater fish, in 1976 and 1977, from
    selected sites throughout the United States, and found mercury levels
    of 0.01-0.84 mg/kg wet weight.

         Berg et al. (1966) analysed feathers from Swedish birds collected
    over a period of 100 years, and found roughly constant levels of
    mercury during the period 1840 to 1940. However, a well documented
    increase of 10-20 times appeared in the 1940s and 1950s, which the
    authors concluded was due to the use of alkylmercury seed dressings.
    Martin (1972) and Martin & Nickerson (1973) sampled starlings
    throughout the United Slates in 1970 and 1971 and found that most of
    the birds had mercury levels of < 0.5 mg/kg (76% of the birds

    analysed in 1971 contained levels of < 0.05 mg/kg). Lindsay & Dimmick
    (1983) found mercury in the liver, breast muscle, and body fat of wood
    duck taken from the area of the Holston River, Tennessee, USA. The
    highest levels were in juveniles (0.42, 0.15, and 0.1 mg/kg, for the
    three tissues, respectively. Local sediment contained
    0.76 mg mercury/kg, black fly larvae and aquatic plants < 0.1 mg/kg.

         Osborn & Nicholson (1984) sampled puffin from the islands of
    St. Kilda and May, off the British coast, and found liver and kidney
    mercury levels of approximately 1.25 mg/kg dry weight (in both
    tissues) for the Isle of May, and 3.75 and 5 mg/kg dry weight,
    respectively, for St. Kilda. Braune (1987) analysed tissues of nine
    species of sea birds sampled in New Brunswick, Canada, for total
    mercury content, and found highest levels in the liver (0.046 to
    0.606 mg/kg) and kidney (0.242 to 5.345 mg/kg). Birds which fed on
    benthic invertebrates or fish had the highest levels, while those
    feeding mainly on pelagic invertebrates had the lowest.

         Fimreite et al. (1982) sampled eggs from a Norwegian gannet
    colony for mercury in 1972, 1978, and 1979, and obtained values of
    0.58, 0.8, and 0.36 mg/kg, respectively. Ohlendorf (1986) analysed
    eggs from three Hawaiian seabird species in 1980, and found mercury in
    all eggs, ranging from 0.122 to 0.359 mg/kg wet weight. Koeman et al.
    (1975) analysed oiled seabirds (guillemot and razorbill) from the
    Dutch coast for mercury residues and reported levels ranging from 1.8
    to 2.4 mg/kg wet weight. Hoffman & Curnow (1979) analysed the levels
    of mercury in the tissues of herons, egrets, and their food collected
    from two sites near Lake Erie, USA. One population fed on Lake Erie
    (food items, 0.02-0.81 mg/kg wet weight; bird livers, 3.0-16.5 mg/kg
    wet weight). The other population fed predominantly on bordering
    marshland (food items, up to 0.24 mg/kg; bird livers,
    1.03-8.22 mg/kg).

         Honda et al. (1986) sampled striped dolphin  (Stenella
     coeruleoalba) and found that the accumulation of total mercury in
    bone correlated significantly with age. Levels rose to 1.44 and
    1.55 mg/kg for adult male and female, respectively, and similar trends
    were seen for methylmercury, levels reaching 0.27 mg/kg in adults.
    Falconer et al. (1983) found that in common porpoise  (Phocoena
     phocoena) highest mercury levels were in the liver, where mean
    levels for females were 6.03 mg/kg and for males 3.42 mg/kg.
    Heppleston & French (1973) analysed tissues of common and grey seals,
    from the British coast for mercury and found highest levels in the
    livers (4.9-113 mg/kg). Koeman et al. (1975) determined mercury levels
    of 0.37-326 mg/kg in the livers of marine mammals (seals, dolphins,
    and porpoises) and also reported an almost perfect correlation between
    mercury and selenium content of these mammals (1:1 ratio between
    mercury and selenium concentrations). The authors suggested that
    selenium uptake may protect marine mammals from the toxic effects of

    mercury. Gaskin et al. (1974) found liver total mercury levels ranging
    from 13 to 157 mg/kg in short-finned pilot whales and long-snouted
    dolphins from the Lesser Antilles. Between 2% and 17% of the total
    mercury was methylated.

    4.4.2  Mercury manufacturing and general industrial areas

         Yeaple (1972) analysed bryophytes from various localities of
    eastern USA for mercury content and found that highest levels
    (1.45 mg/kg) were in plants from a large city. Levels in cities and
    industrial areas were higher than those in rural areas (e.g.,
    < 0.05 mg/kg in a high, isolated mountain area). Kraus et al. (1986)
    collected leaves of the salt marsh cordgrass  (Spartina alterniflora)
    from two sites in the USA, one site near a heavily industrialized area
    and the other in a non-industrialized area. The mean soil
    concentrations of mercury for the two sites were 18.17 and 0.22 mg/kg,
    respectively, while the residues in the leaves were 0.16 and
    0.02 mg/kg, respectively. Salts collected from the surface of plants
    in the contaminated area contained 0.11 mg mercury/kg; laboratory
    studies have shown the plant capable of mercury excretion.

         Nuorteva et al. (1980) analysed trout  (Salmo trutta) from the
    Idrijca River, Yugoslavia, about 3 km downstream from a mercury
    distillation plant. The fish had a mercury content of 0.66 mg/kg in
    the flesh, and highest levels were found in the spleen and kidney
    (17.5 and 24 mg/kg, respectively). Three samples of ephemerids, taken
    6 km from the plant, contained 0.27, 0.36, and 0.56 mg/kg wet weight,
    and a sample containing 4.28 mg/kg was found 1 km from the plant.
    These were lower levels than those reported previously, presumably
    because of six months inactivity at the plant. The same authors
    analysed blow flies from various polluted and non-polluted localities.
    From an unpolluted area mercury levels were < 0.1 mg/kg, near a
    Finnish pulp factory, 0.2 mg/kg, and near a caustic soda factory,
    0.3 mg/kg. Higher levels (0.8 mg/kg) were found close to a mercury
    mine and distillation plant in Yugoslavia, whereas levels were near
    normal 1 km upstream or downstream from the mine.

         Doi et al. (1984) analysed feathers from birds collected over a
    period of 25 years from the mercury-polluted shores of the Shiranui
    Sea, Japan. Relatively high levels were found until the late 1970s
    even though the draining of water containing methylmercury from a
    local factory was stopped in 1968. Mean mercury levels were: fish-
    eating birds, 7.1 mg/kg; omnivorous waterfowl, 5.5 mg/kg; predatory
    birds, 3.6 mg/kg; omnivorous terrestrial birds, 1.5 mg/kg; and
    herbivorous waterfowl, 0.9 mg/kg.

         Fimreite et al. (1971) analysed 156 fish and 48 bird livers from
    the Great Lakes area of Canada in 1968 and 1969. Elevated mercury
    levels were found in all fish samples, highest levels occurring in
    lake trout, pumpkinseed sunfish, and walleye (10.5, 7.09, and

    5.01 mg/kg, respectively). Levels were generally highest in fish
    collected downstream from suspected sources. The highest mercury level
    in a fish-eating bird was found in a red-necked grebe, where the liver
    level was 17.4 mg/kg. Three grebes sampled showed a range of
    0.45-17.4 mg/kg. Lower concentrations were found in cormorants,
    herons, murrelets, terns, kingfisher, and other fish-eating birds, but
    mean mercury liver burden was greater in these birds than in non fish-
    eating species.

    4.4.3  Mining activity

         Huckabee et al. (1983) monitored levels of mercury in vegetation
    in the vicinity of the mercury mine at Almaden in Spain. Mean
    concentrations of total mercury in vegetation ranged from > 100 mg/kg
    within 0.5 km of the mine to 0.20 mg/kg 20 km from the mine. There was
    still a significantly higher mercury content in vegetation 25 km
    upwind from the mine (about 10 times the background level). Mosses
    were found to contain the greatest concentration of mercury
    (7.58 mg/kg), and woody plants accumulated less of the metal
    (0.72 mg/kg) than herbaceous plants (2.25 mg/kg). The figures given
    are for samples collected in spring. There was a correlation between
    distance from the mine and plant mercury content for woody plants and
    mosses but not for herbaceous plants. No methylmercury, at
    quantifiable levels, was found in any of the plants analysed, although
    traces were seen in several samples indicating a methylmercury content
    of less than 10 pg per sample.

         When Phillips & Buhler (1980) analysed rainbow trout  (Salmo
     gairdneri), stocked in a reservoir contaminated by a disused mine,
    for mercury, they found that lateral muscle tissue levels increased
    linearly during the first five months that the fish were in the
    reservoir. Trout sampled 7, 19, or 31 months after being introduced
    showed levels that did not differ significantly (mean level = 1.25 mg
    mercury/kg). Matsunaga (1975) analysed crucian carp, dace, and zacco
    temmincku from two rivers receiving discharge from mercury mines in
    Japan. Total mercury levels in the fish were approximately
    0.2-4.5 mg/kg and reflected the levels of mercury in the water
    (4-50 ng/kg).

         Hesse et al. (1975) determined total mercury concentrations in
    the muscle, liver, and kidney of 22 species of birds collected from a
    western South Dakota watershed contaminated by mining activity.
    Elevated mercury levels were found in fish-eating birds, especially
    double-crested cormorants. Levels in non fish-eating birds were lower
    but still significantly higher than background. In general, greater
    accumulations occurred in the livers of fish-eating birds (0.89 to
    30.9 mg/kg) and in the kidneys of non-fish-eating birds (0.27 to
    0.60 mg/kg).

    4.4.4  Chloralkali plants

         Gardner et al. (1978) analysed sediment, plants, and animals from
    a salt marsh contaminated by a chloralkali plant in Brunswick,
    Georgia, USA. Chloralkali plants produce metallic mercury from salts.
    Sediment levels ranged from 0.27 to 1.7 mg/kg dry weight for the top
    5 cm and they varied according to distance from plant and depth of
    sample. The roots of  Spartina alterniflora, the marsh grass,
    contained the highest levels (0.07-1.47 mg/kg dry weight) within the
    plant. Of the animals analysed from the contaminated marsh and nearby
    river, the invertebrates contained 0.3-9.4 mg/kg dry weight, the fish
    0.3-1.9 mg/kg dry weight, the birds 2.4-37.0 mg/kg dry weight (liver)
    and the mammals 3.8-15 mg/kg dry weight (liver). Methylmercury levels
    were low (< 0.002 mg/kg) in sediment and plants but accounted for
    most of the mercury found in the tissues of higher organisms.

         Hildebrand et al. (1980) sampled fish and invertebrates from the
    Holston River, USA, above and below an inactive chloralkali plant.
    Rock bass and hog sucker contained total mercury levels at less than
    1 mg/kg above the plant, and 1-3 mg/kg immediately below it. Benthic
    invertebrates gave a similar pattern, lower levels being found above
    the plant and the higher levels below it. Total mercury concentrations
    in the individual taxonomic groups of the invertebrates ranged from a
    maximum of 3.75 mg/kg  (Hydropsychidae, 3.7 km below the plant) to a
    minimum of 0.016 mg/kg  (Psepheridae, 5.5 km above the plant). Total
    mercury concentrations in fish and invertebrates decreased with
    distance down stream of the plant. Mercury in the methyl form
    comprised 91.7% of total mercury in the fish and 50% in the
    invertebrates.

         Wallin (1976) reported that samples of the carpet-forming moss
     Hypnum cupressiforme from sites around six Swedish chloralkali
    plants all contained similar mercury levels. Levels were highest
    (1-15 mg/kg) close to the plants and decreased with increasing
    distance from each plant. Background levels for the region
    (90-150 µg/kg) were reached at distances of 9-15 km from the plants.
    The author calculated that only a small part of the annual fallout
    (< 10%) was deposited locally. Shaw & Panigrahi (1986) analysed soil
    and five species of dwarf plants, from an area adjacent to a
    chloralkali factory, for mercury content. Soil from around the roots
    of the plants was analysed, and the mercury content was found to be
    very variable (2.13-893 mg/kg dry weight). Uptake into the roots,
    stem, leaf, and fruit of all plants in the area was significant.
    Leaves contained the highest levels of mercury, ranging between 2.32
    and 38.8 mg/kg dry weight. Greater accumulation of mercury was found
    in the stem than roots of  Croton sp. and  Jatropha sp.; similar
    amounts in both stem and roots of  Argemone sp., and more mercury in
    the roots than the stem of  Ipomoea sp. and  Calotropis sp. No
    correlation was found between the soil mercury level and plant uptake.
    Bull et al. (1977) measured mercury in soil, grass, earthworms, and

    small mammals near a chloralkali factory. At a distance of < 0.5 km
    from the factory, mean mercury levels in surface soil (3.81 mg/kg dry
    weight), grass (4.01 mg/kg dry weight), earthworms (1.29 mg/kg wet
    weight) and moss bags (63 ng/dm2 per day) were significantly higher
    than levels found 10 to 30 km from the works. Levels of mercury at
    this distance were comparable with those found at sites not associated
    with mercury sources. Mercury levels in all tissues analysed, except
    muscle of bank voles  (Clethrionomys glareolus) and woodmice
     (Apodemus sylvaticus) were significantly higher in the study area
    than control areas. The authors also found elevated levels of
    methylmercury in small mammals and earthworms in the study area,
    suggesting methylation of the inorganic mercury fall-out.

    4.4.5  Mercurial fungicides

         Fimreite et al. (1970) found that seed-eating birds, and their
    avian predators, had higher liver mercury levels in areas where
    treated grain (mercurial fungicide) had been sown compared with areas
    using untreated grain. Jefferies & French (1976) analysed specimens of
    the long-tailed field mouse  (Apodemus sylvaticus) taken from a wheat
    field that had been drilled two months previously with wheat dressed
    with dieldrin and mercury. Whole body mercury concentrations were much
    higher (0.83 ± 0.44 mg/kg wet weight) than those found immediately
    after drilling (0.39 ± 0.04 mg/kg wet weight).

    5.  TOXICITY TO MICROORGANISMS

         Mercury in an inorganic form is toxic to microorganisms. It is
    much more toxic in an organic form, owing to increased availability of
    the metal to cells. The following are illustrative examples, rather
    than an exhaustive cover, of research into the effects of mercury on
    microorganisms.

         Wood (1984) discussed six protective mechanisms available to
    microorganisms (and certain higher organisms) that increase their
    resistance to metal ions in general, and specifically to mercury.
    These mechanisms are biochemical in nature and, generally, render the
    mercury ion ineffective in disturbing the normal biochemical processes
    of the cell. The mechanisms are: (a) efflux pumps that remove the ion
    from the cell, a process which requires energy, (b) enzymatic
    reduction to the less toxic elemental form; (c) chelation by
    intracellular polymers (not firmly established for mercury);
    (d) binding of mercury to cell surfaces; (e) precipitation of
    insoluble inorganic complexes, usually sulfides and oxides, at the
    cell surface; and (f) biomethylation with subsequent transport through
    the cell membrane by simple diffusion. It is this last mechanism,
    biomethylation, which renders the mercury more toxic to higher life-
    forms.

    5.1  Toxicity of Inorganic Mercury

    Appraisal

          Inorganic mercury is toxic to microorganisms over a wide range
     of concentrations. Its effects on development and survival are
     modified by environmental factors such as temperature, light
     intensity, pH, and chemical composition of the medium, and by cell-
     related factors such as genetic variation. Through selective effects
     on particular species, it can change the composition of a plankton
     community. The mechanism of action is not fully understood.

    5.1.1  Single species cultures

         Kamp-Nielson (1971) demonstrated a time-dependent effect of
    mercuric chloride, added at 300 µg/litre, on the photosynthesis of
     Chlorella pyrenoidosa. There was little effect in the first hour of
    incubation, a pronounced drop in photosynthetic rate in the second
    hour, and a period of little further effect between 2 and 5 h. An
    overall rate reduction of about 50% occurred after 5 h with a cell
    density of 6.5 × 107 cells/litre. There was a greater effect on
    photosynthesis at lower cell densities. It was also found that
    photosynthesis had to occur for the effect to develop, since exposure
    to mercuric chloride for 2 h in the light had the same effect as
    exposure to the same concentration of mercury for 2 h in the dark
    followed by 2 h in the light. Similar results were found after 1-h
    exposures in light and darkness followed by light. There was an effect

    of light intensity; in short-term experiments mercury had a
    deleterious effect on photosynthesis only at high light intensities.
    Mercury also affected photosynthesis at low light intensity, but only
    after 20-h exposures. Mercury affected photosynthesis adversely at
    concentrations between about 50 and 300 µg/litre, but had no greater
    effect at concentrations up to 1000 µg/litre (the highest tested). The
    effect was dependant on cell density, pH, light intensity, and
    duration of exposure. Potassium and sodium in the growth medium had no
    effect on mercury toxicity to  Chlorella. Increasing the
    concentration of mercuric chloride in the medium increased the
    "leakage" of potassium from the cells of  Chlorella. This was maximal
    at a mercury concentration of about 300 µg/litre and was considered to
    be the main toxic effect of mercury. The effect on potassium leakage
    occurred equally in darkness and light and was, therefore, independent
    of the photosynthetic effect. Mercury increased the length of the 
    lag-phase during the growth of  Chlorella pyrenoidosa cultures. A
    greater effect was seen at 660 than at 330 µg/litre, the only two doses
    tested. This effect was also demonstrated by Osokina et al. (1984) in
    the green alga  Scenedesmus quadricauda. The effect was highly
    dependant on the cell density of the original inoculum.

         Rai et al. (1981) exposed  Chlorella vulgaris to mercuric
    chloride concentrations between 100 and 1000 µg/litre for 3 weeks, and
    monitored growth and survival. LC50 for survival was at 400 µg/litre
    of mercuric chloride. The growth rate was 92% of the control value at
    100 µg/litre and 31% at 800 µg/litre, and there was no growth at
    1000 µg mercuric chloride/litre. The chlorophyll content of the cells
    was reduced throughout the dose range. There was a greater toxic
    effect of mercuric chloride at low pH, with the greatest amelioration
    of toxicity at pH 9. There was also a protective effect of calcium and
    phosphate in the medium and, to a lesser extent, of magnesium. Both
    calcium and phosphate increased the yield of algae, in the presence of
    sublethal concentrations of mercury, when added at concentrations up
    to 20 mg/litre. At higher concentrations of both calcium and
    phosphate, the protection was less marked. Den Dooren de Jong (1965)
    determined the no-observed-effect-level (NOEL) for mercuric chloride
    on  Chlorella vulgaris to be 50 µg/litre. Hannon & Patouillet (1972)
    emphasized the irreversibility of the effects of mercuric chloride on
     Chlorella pyrenoidosa. If mercury was present in sufficient
    concentration to affect growth of the alga, then no recovery was found
    following transfer in clean medium. Similar effects were reported for
    three species of marine unicellular algae. Mercury toxicity was
    dependant on cell numbers in the initial inoculum (Kuiper, 1981). In
    studies with unialgal cultures of  Chlamydomonas sp., there was a
    relationship between cell concentration and mercury toxicity. The
    author attributed this to a surface area effect, the metal is being
    adsorbed onto cell walls to cause its effect on the unicellular algae.

         Huisman et al. (1980) investigated the effect of temperature on
    the toxicity of mercuric salts to the green alga  Scenedesmus acutus.
    Mercury concentration in the cultures was kept constant by a

    mercury(II) buffer system, and the growth and photosynthesis of the
    alga were monitored. Toxicity increased with increasing temperature
    over the range 15-30°C. There was no effect observed in this study on
    the lag phase, no later increase in growth, and no effect of initial
    cell numbers. This was attributed to the buffer system which prevented
    changes in free mercury concentrations over time. The authors also
    examined the binding of mercury to algal cells. Metal bound to the
    cell wall consists of two fractions: one which can be washed off with
    cysteine solution and one which cannot. The amount of mercury which
    can be washed off the cell wall increase with increasing temperature.
    The mercury bound to cell walls, but washable with cysteine, appears
    to be the toxic fraction. The total mercury content of algal cells
    does not correlate with effect. A total mercury content not lethal at
    15°C causes complete inhibition of growth and photosynthesis at 30°C.
    Recovery occurs under circumstances where the cells retain the non-
    washable mercury, indicating that the washable fraction is the toxic
    component. The authors suggested that the reversibility of the action
    of cysteine-washable mercury indicates that the metal is bound to
    carboxyl or phosphate groups and not to sulfhydryl groups. These
    mercury ions can be readily exchanged for other metal ions, leading to
    a decreased inhibition by mercury. Therefore, in media with a high
    concentration of dissolved salts, mercury appears to be less toxic.
    The authors postulated another mechanism by which mercury might be
    toxic to algal cells. Interference with potassium-sodium-dependent
    ATPase in the cell membrane influences the active transport of
    nutrients. This would give rise to disturbances of nitrogen metabolism
    and also of photosynthesis. The delayed action of mercury on cultures
    could be ascribed to their being initially rich in nitrogen, and,
    therefore, less susceptible to nitrogen starvation.

         Nuzzi (1972) exposed  Phaeodactylum tricornutum, Chlorella sp.,
    and  Chlamydomonas sp., isolated from the lower Hudson River, New
    York, USA, to mercuric chloride. The growth of all three organisms was
    severely inhibited by mercury at 7.5 µg/litre (to between 50% and 75%
    of control growth). The growth of  Chlamydomonas sp. was completely
    inhibited by 15 µg mercuric chloride/litre and the other two species
    by 22 µg/litre.

         Gray & Ventilla (1971) found no effect of mercuric chloride on
    growth of