The secondary active Clsecretion in seawater (SW) teleost fish gills and elasmobranch rectal gland involves basolateral Na + ,K + -ATPase and NKCC, apical membrane CFTR anion channels, and a paracellular Na + -selective conductance. In freshwater (FW) teleost gill, the mechanism of NaCl uptake is more controversial and involves apical V-type H + -ATPase linked to an apical Na + channel, apical Cl � -HCO3 exchange and basolateral Na + ,K + -ATPase. Ca 2+ uptake (in FW and SW) is via Ca 2+ channels in the apical membrane and Ca 2+ -ATPase in the basolateral membrane. Mainly this transport occurs in mitochondria rich (MR) chloride cells, but there is a role for the pavement cells also. Future research will likely expand in two major directions, molded by methodology: first in physiological genomics of all the transporters, including their expression, trafficking, operation, and regulation at the molecular level, and second in biotelemetry to examine multivariable components in behavioral physiological ecology, thus widening the integration of physiology from the molecular to the environmental levels while deepening understanding at all levels. J. Exp. Zool. 293:264-283, 2002. r 2002 Wiley-Liss, Inc.

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Na+, Cl−, Ca2+ and Zn2+ transport by fish gills: retrospective review and prospective synthesis

There is a long research history addressing olfactory imprinting, natal homing, and non-natal straying by anadromous salmon and trout (Salmoni- dae). In undisturbed populations, adult straying is a fundamental component of metapopulation biology, facilitating genetic resilience, demographic stability, recolonization, and range expansion into unexploited habitats. Unfortunately, salmonid hatcheries and other human actions worldwide have affected straying in ways that can negatively affect wild populations through competitive interactions, reduced productiv- ity and resiliency, hybridization and domestication effects, and outbreeding depression. Reduced adult straying is therefore an objective for many managed populations. Currently, there is considerable uncer- tainty about the range of 'natural' stray rates and about which mechanisms precipitate straying in either wild or human-influenced fish. Research in several disci- plines indicates that adult straying is affected by endocrine physiology and neurological processes in juveniles, incomplete or interrupted imprinting during rearing and emigration, and by complex interactions among adult maturation processes, reproductive behaviors, olfactory memory, environmental condi- tions during migration, and senescence physiology. Reported salmonid stray rates indicate that the behavior varies among species, among life-history types, and among populations within species. Most strays enter sites near natal areas, but long-distance straying also occurs, especially in hatchery popula- tions that were outplanted or transported as juveniles. A majority of past studies has estimated straying as demographic losses from donor populations, but some have estimated straying into recipient populations. Most recipient-based estimates have substantiated concerns that wild populations are vulnerable to swamping by abundant hatchery and farm-raised strays.

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Homing and straying by anadromous salmonids: a review of mechanisms and rates

We review factors affecting the within-river spawning migration of Atlantic salmon. With populations declining across the entire distribution range, it is important that spawners survive in the last phase of the spawning migration. Knowledge on the factors affecting migration is essential for the protection of populations, and to increase the success of reintroduction programmes. A number of studies have documented that the upstream migration may be delayed for many weeks at man-made obstacles such as power station outlets, residual flow stretches, dams, weirs and fishways. The fish may also be delayed at natural migration barriers. Often, the magnitude of delay is not predictable; fish may be considerably delayed at barriers that appear to humans to be easily passable, or they may quickly pass barriers that appear difficult. Stressful events like catch-and-release angling may affect upstream migration. Impacts of human activities may also cause altered migration patterns, affect the within-river distribution of the spawning population, and severe barriers may result in displacement of the spawning population to other rivers. Factors documented to affect within-river migration include previous experience, water discharge, water temperature, water velocity, required jump heights, fish size, fish acclimatisation, light, water quality/pollution, time of the season, and catch and handling stress. How each of these factors affects the upstream migration is to a varying extent understood; however, the effects may differ among different river sections and sites. There are likely a number of additional important factors, and the relationship between different factors is complex. The understanding of general mechanisms stimulating fish within-river migration are still lacking, and it cannot be reliably predicted under which conditions a fish will pass a given migration barrier or which conditions are needed to stimulate migration at different sites. The strong focus on the effects of water discharge in past work may have hampered consideration of other factors. Exploration of the influence of these other factors in future studies could improve our understanding of what controls the upstream migration.

Factors affecting the within-river spawning migration of Atlantic salmon, with emphasis on human impacts

Electromyogram (EMG) telemetry studies that involve remotely monitoring the locomotory activity and energetics of fish are contributing important information to the conservation and management of fisheries resources. Here, we outline the development of this rapidly evolving field and formulate the studies conducted that utilize this technology. To date, more than 60 studies have been conducted using EMG telemetry that spans 18 species. Several general trends were observed in the methodology of the studies that we have highlighted as standards that should be adopted associated with transmitter customization, electrode placement and surgical technique. Although numerous studies have been methodological, there are still some deficiencies in our basic understanding of issues such as the need for individual calibration and the method of reporting or transforming data. Increasingly, this technology is being applied to address issues in conservation, management and aquaculture production. At present, the technology has been most frequently applied to the study of animal activity or energetics and to migration. Several recent studies have also focused on addressing more basic questions in ecological and evolutionary biology (e.g. parental care dynamics) similar to the large body of literature that has been collected for other taxa (e.g. marine mammals, birds), using activity telemetry. Collectively, studies conducted using EMG telemetry have contributed important information on free-swimming fish that was previously difficult to obtain. EMG telemetry is particularly effective for examining behaviour at temporal and spatial scales that are difficult using other techniques. The development of an ultrasonic transmitter based on the same proven principles as those used in the current radio transmitter technology will permit studies in other environments (i.e. marine, brackish, deep water) and on different species of fish. We encourage the continued development and refinement of devices for monitoring the activity and energetics of free-swimming fish, and also encourage researchers to consider EMG telemetry as a tool for addressing questions that are not effectively answered with other techniques.

Activity and energetics of free-swimming fish: insights from electromyogram telemetry

Chemical reactions that involve radical intermediates can be influenced by mag- netic fields, which act to alter their rate, yield, or product distribution. These effects have been studied extensively in liquids, solids, and constrained media such as micelles. They may be interpreted using the radical pair mechanism (RPM). Such effects are central to the field of spin chemistry of which there have been several detailed and extensive reviews. This re- view instead presents an introductory account of the field of spin chemistry, suitable for use by graduate students or researchers who are new to the area. It proceeds by giving a brief his- torical overview of the development of spin chemistry, before introducing the essential the- ory. This is then illustrated by application to a series of recent developments in solution-phase magnetic field effects (MFEs). The closing pages of this review describe the role played by spin chemistry in the remarkable magnetic compass sense of birds and other animals.

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Magnetic field effects in chemical systems

From a total of 174 multi-sea-winter Atlantic salmon radio tagged in the Tanafjord (northern Norway, 70° N) during 1992 and 1993, 48 Atlantic salmon were followed from entering the River Tana until spawning. Three phases were identified: (1) migratory, direct or stepwise migration to, or close to the position held at spawning; (2) search, movements both up and down river at or close to the position held at spawning; (3) holding, a period without movements prior to spawning. During the migratory phase, Atlantic salmon migrated directly to near the spawning area, or stopped between one and nine shorter periods during the upstream migration. Number of stops increased with increasing migratory distance in 1993, but no such correlation was found in 1992. The highest migratory speeds were recorded in the lower parts of the river. A distinct change in migratory pattern was found in 67% of the Atlantic salmon near or at the area held at spawning. Most common was a search phase of erratic movements with more than one down river movement. After the movement terminated, 96% of the Atlantic salmon had a period when no or little movement was recorded until spawning (on average 55 days in 1992 and 51 days in 1993). There was no preference for staying at, up or down river from the spawning area during this holding period. Early ascending Atlantic salmon migrated to spawning areas further from the mouth than the later arriving Atlantic salmon in 1993, but not in 1992. The proportion of time spent on the migratory phase increased, while the proportion of time spent on the holding phase decreased with increasing distance to the spawning area.

Return migration of Atlantic Salmon in the River Tana: Phases of migratory behaviour

Swimming speed and average electromyogram (EMG) pulse intervals were highly correlated in individual lake trout Salvelinus namaycush (r 2 =0.52-0.89) and brown trout Salmo trutta (r 2 =0.45-0.96). High correlations were found also for pooled data in both lake trout (r 2 =0.90) and brown trout of the Ema stock (r 2 =0.96) and Laerdal stock (r 2 =0.96). The linear relationship between swimming speed and average EMG pulse intervals differed significantly among lake trout and the brown trout stocks. This successful calibration of EMGs to swimming speed opens the possibility of recording swimming speed of free swimming lake trout and brown trout in situ. EMGs can also be calibrated to oxygen consumption to record energy expenditure.

Radio-transmitted electromyogram signals as indicators of swimming speed in lake trout and brown trout

American eels Anguilla rostrata, collected from three distinct locations along the St Lawrence River (Lake St Lawrence, Quebec City and Kamouraska), were thought to consist entirely of out-migrating adults at Quebec City and Kamouraska and of both resident and migrants at Lake St Lawrence. The mean plasma levels of 17β-oestradiol closely paralleled the trend observed with gonadosomatic index (I G ) and oocyte development. The highest levels of 17β-oestradiol and I G were observed in Quebec City and Kamouraska, while the levels observed in the resident eels from Lake St Lawrence were very low. Generally, the concentration of total plasma non-esterified fatty acids (NEFA) appeared to follow the trend of 17β-oestradiol and I G and increased with sexual maturity. However, the most sexually mature group, eels from Kamouraska, had the lowest total plasma NEFA levels. The reason for the drop is not clear at present. In addition, a number of shifts in the plasma NEFA profiles of American eels did occur during their migration in a downstream direction, with several fatty acids either increasing or decreasing in relative abundance. The ratio of n3: n6 fatty acids declined with increasing sexual maturity as most n3 fatty acids became less abundant. Although the correlation between gonadal development and the relative concentration of individual fatty acids was not always clear, both arachidonic acid (20: 4n6) and docosahexaenoic acid (22: 6n3) were significantly higher in the plasma of out-migrating adults than in resident yellow phase eels.

R.S. McKinley

Papers

Return migration of Atlantic Salmon in the River Tana: Phases of migratory behaviour

Radio-transmitted electromyogram signals as indicators of swimming speed in lake trout and brown trout

Plasma non-esterified fatty acid profiles and 17β-oestradiol levels of juvenile immature and maturing adult American eels in the St Lawrence River