<br />60 HAMILTON ET AL
<br />
<br />or intermittently to 30 /-Lg/L of copper for 56 days
<br />postfertilization (30 days pre hatch and 26 days
<br />posthatch), whereas survival was reduced at 51 /-Lg/L
<br />after 38 days in the intermittent exposure and at 57
<br />/-Lg/L after 63 days in the continuous exposure. How-
<br />ever, effects in their study occurred at lower whole-body
<br />copper residues (6-10 /-Lg/g) than in our study (23
<br />/-Lg/g and higher). Buckley et al. (1982) also reported
<br />reduced growth in coho salmon (Oncorhynchus kisutch)
<br />exposed to 70 /-Lg/L of copper for as little as 2 weeks,
<br />but no effect on survival even in fish exposed to 140
<br />/-Lg/L for 16 weeks. In contrast, Marr et al. (1996)
<br />reported reduced growth in rainbow trout exposed in
<br />soft water (hardness 25 mg/L as CaC03) to 4.6 /-Lg/L
<br />of copper in 2 weeks, but no apparent effects on
<br />survival after 60 days at the highest concentration
<br />tested, 9 /-Lg/L.
<br />Exposure to zinc concentrations of 79 /-Lg/L in the
<br />razorback sucker study (4X) and 130 /-Lg/L in the
<br />bonytail study (8X) also could have contributed to
<br />reduced growth and survival of fish. Bengtsson (1974b)
<br />reported reduced growth in the minnow Phoxinus phox-
<br />inus exposed in soft water (hardness 39 mg/L as
<br />CaC03) for 30 days to 130 p.g/L of zinc, whereas in a
<br />separate experiment with the same species tested in
<br />similar water (hardness 47 mg/L as CaC03) survival
<br />was reduced at 80 p.g/L (Bengtsson, 1974c). Spehar
<br />(1976) reported that female flagfish (Jordanella flori-
<br />dae) exposed in soft water (hardness 44 mg/L as
<br />CaC03) to zinc had reduced growth at 51 /-Lg/L after
<br />100 days of exposure and reduced survival at 85 /-Lg/L
<br />at 30 days. Sinley et al. (1974) reported increased
<br />mortality in swim-up rainbow trout exposed in soft
<br />water (25 mg/L as CaC03) to 260 /-Lg/L of zinc.
<br />Concentrations of copper, selenium, and zinc that
<br />cause adverse effects discussed above fall within the
<br />range of concentrations tested in the present study, and
<br />thus, suggest that they caused the adverse effects ob-
<br />served. Copper and zinc are essential inorganics neces-
<br />sary to meet the metabolic needs of animals, and are
<br />homeostatic ally controlled in fish by metallothionein, a
<br />protein found in liver and kidney tissue (Hamilton and
<br />Mehrle, 1986). Metallothionein also serves a detoxifica-
<br />tion role by sequestering inorganics such as cadmium
<br />and mercury, as well as excessive amounts of copper
<br />and zinc, and reducing the amount of free metals in
<br />tissues, thereby reducing potential toxicity.
<br />Selenium is also an essential micronutrient to ani-
<br />mals as a component of certain proteins and enzymes
<br />where it is selectively incorporated against a concentra-
<br />tion gradient of sulfur, which is present at concentra-
<br />tions that are orders of magnitude higher than sele-
<br />nium concentrations (Stadtman, 1980). However, when
<br />selenium is available at greater than micro molar con-
<br />centrations, it can be extensively substituted indiscrimi-
<br />
<br />nately for sulfur in many cellular constituents (Stadt-
<br />man, 1974). Although there is a mechanism for the
<br />metabolic control of copper and zinc, i.e., metallqth-
<br />ionein, there is no known control for selenium. Stadt-
<br />man (1974) stated "because of the greater reactivity
<br />and lower stability of selenium compounds compared
<br />to the corresponding sulfur compounds, the cell may
<br />encounter metabolic problems which eventually can
<br />lead to death of the organism." The death of fish and
<br />wildlife from selenium poisoning has been documented
<br />in 12 case histories by Skorupa (1998).
<br />The consequence of reductions in total length,
<br />weight, or condition factor is that swimming perfor-
<br />mance is reduced, and susceptibility to predation is
<br />increased (Barns, 1967; Taylor and McPhail, 1985), and
<br />feeding efficiency is reduced (Gunn and Noakes, 1987).
<br />Functions that promote fish survival are also related to
<br />body size (reviewed by Ojanguren et aI., 1996). For
<br />example, sensory systems develop rapidly during e"r1y
<br />life stages and visual acuity increases with fish siZe,
<br />enabling large fish to better detect predators and fqod
<br />and resist starvation, thus allowing them access to a
<br />wider range of prey, which increases the likelihood of
<br />finding appropriate food. Ojanguren et al. (1996) fur-
<br />ther stated that better swimming performance by larger
<br />fish could reflect the overall fitness because swimming
<br />is a highly integrative function requiring muscular
<br />strength and coordination, morphometric adequacy, ac-
<br />curate sensorial perception, and physiological adjl.\st-
<br />ments.
<br />It has been hypothesized that interaction of slOw
<br />growth and increased early-life mortality have con-
<br />tributed to the decline of the endangered Colorado
<br />squawfish in the upper Colorado River (Kaeding and
<br />Osmundson, 1988). It is also believed that the upper
<br />Colorado River provided suboptimal conditions for fish
<br />growth because of generally lower water temperatu~es
<br />compared to the lower basin. Thompson et a!. (1990
<br />assessed this hypothesis and found that smaller Col-
<br />orado squawfish with lower condition factors had ~e-
<br />duced survival during overwinter conditions compar~d
<br />to larger fish with higher condition factors. Others have
<br />also shown that overwinter survival of a variety of otlier
<br />fish species depends on fish size and condition factor
<br />(reviewed by Thompson et aI., 1991). Overwinter sur-
<br />vival of fish has also been shown to be reduced by
<br />exposure to selenium at low concentrations in water
<br />(4.8 /-Lg/L) and diet (5.1 p.g/g) (Lemly, 1993a). In t~at
<br />study, stress from low water temperatures and selenium
<br />residues caused hematological changes and gill damage
<br />that reduced respiratory capacity, reduced activity and
<br />feeding, depleted body lipids by 50-80%, and reduced
<br />survival. Consequently, razorback sucker and bony tail
<br />with reduced growth may have reduced survival poten-
<br />tial. .
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