Mismatch between fishway operation and timing of fish movements: a risk for cascading effects in partial migration systems

Habitat fragmentation is a growing problem worldwide. Particularly in river systems, numerous dams and weirs hamper the movement of a wide variety of species. With the aim to preserve connectivity for fish, many barriers in river systems are equipped with fishways (also called fish passages or fish ladders). However, few fishways provide full connectivity. Here we hypothesized that restricted seasonal opening times of fishways can importantly reduce their effectiveness by interfering with the timing of fish migration, for both spring‐ and autumn‐spawning species. We empirically tested our hypothesis, and discuss the possible eco‐evolutionary consequences of affected migration timing. We analyzed movements of two salmonid fishes, spring‐spawning European grayling (Thymallus thymallus) and autumn‐spawning brown trout (Salmo trutta), in Norway's two largest river systems. We compared their timing of upstream passage through four fishways collected over 28 years with the timing of fish movements in unfragmented river sections as monitored by radiotelemetry. Confirming our hypothesis, late opening of fishways delayed the migration of European grayling in spring, and early closure of fishways blocked migration for brown trout on their way to spawning locations during late autumn. We show in a theoretical framework how restricted opening times of fishways can induce shifts from migratory to resident behavior in potamodromous partial migration systems, and propose that this can induce density‐dependent effects among fish accumulating in lower regions of rivers. Hence, fragmentation may not only directly affect the migratory individuals in the population, but may also have effects that cascade downstream and alter circumstances for resident fish. Fishway functionality is inadequate if there is a mismatch between natural fish movements and fishway opening times in the same river system, with ecological and possibly evolutionary consequences for fish populations.     

Olfactory gene expression in migrating adult sockeye salmon Oncorhynchus nerka

Expression of 12 olfactory genes was analysed in adult sockeye salmon Oncorhynchus nerka nearing spawning grounds and O. nerka that had strayed from their natal migration route. Variation was found in six of these genes, all of which were olfc olfactory receptors and had lower expression levels in salmon nearing spawning grounds. The results may reflect decreased sensitivity to natal water olfactory cues as these fish are no longer seeking the correct migratory route. The expression of olfactory genes during the olfactory‐mediated spawning migration of Pacific salmon Oncorhynchus spp. is largely unexplored and these findings demonstrate a link between migratory behaviours and olfactory plasticity that provides a basis for future molecular research on salmon homing.     

Importance of the olfactory sense to migratory sea lampreys Petromyzon marinus seeking riverine spawning habitat

This study tested the hypothesis that the sea lamprey Petromyzon marinus, a diadromous species of fish, relies on innately discerned odours, including pheromones, to locate riverine spawning habitat. Migratory, sexually immature P. marinus were captured as they entered streams flowing into the Great Lakes, and their olfactory systems were occluded or not by injecting either innocuous dental impression material or a saline control into their nasopores. Animals were then released back into lakes or streams and their recapture rates in stream traps noted. When released into Lake Huron, P. marinus with intact (functional) olfactory systems were very successful in locating rivers (recapture rates ranged up to 65%), while animals with occluded nasopores were virtually unable to do so and had recapture rates five to 20 times lower than intact animals. With few exceptions, intact fish entered the stream closest to their release point within a few days, irrespective of where they had been originally captured; their ability to locate streams is apparently innate and well developed. In contrast, when released within streams, both intact and occluded P. marinus successfully swam upstream to traps for several days although the ability of the former exceeded that of the latter after this period. Migratory P. marinus rely heavily on olfactory cues, of which a larval pheromone is presumably one, to locate river mouths and to a lesser extent to promote upstream movement within rivers.     

Mechanosensory‐based orientation to elevated prey by a benthic fish

Lake Michigan mottled sculpin, Cottus bairdi, respond to both live and artificial (e.g., vibrating sphere) prey with an unconditioned movement towards the source of vibration, followed by a step‐by‐step approach and final strike at the source. In addition to these well‐studied, whole‐body movements along the horizontal plane of the substrate, sculpin exhibit a little‐studied behavior in which the vertical position of the fish's head can vary from being flush with the substrate to several cm's above the substrate. To test the hypothesis that sculpin can determine source elevation via mechanosensory cues, we measured head elevation of blinded fish as a function of source elevation and distance as fish approached a small (3 mm radius), 50 Hz vibrating sphere. At distances associated with pre‐strike positions (< 2 cm), head elevations were positively correlated with source elevation before but not after pharmacological blocking of the lateral line with CoCl₂. These results demonstrate that sculpin are able to determine source elevation using mechanosensory cues alone and that in the absence of visual and olfactory cues, vertical orientation of the head towards the source requires the lateral line system.     

The Olfactory Transcriptome and Progression of Sexual Maturation in Homing Chum Salmon Oncorhynchus keta

Reproductive homing migration of salmonids requires accurate interaction between the reception of external olfactory cues for navigation to the spawning grounds and the regulation of sexual maturation processes. This study aimed at providing insights into the hypothesized functional link between olfactory sensing of the spawning ground and final sexual maturation. We have therefore assessed the presence and expression levels of olfactory genes by RNA sequencing (RNAseq) of the olfactory rosettes in homing chum salmon Oncorhynchus keta Walbaum from the coastal sea to 75 km upstream the rivers at the pre-spawning ground. The progression of sexual maturation along the brain-pituitary-gonadal axis was assessed through determination of plasma steroid levels by time-resolved fluoroimmunoassays (TR-FIA), pituitary gonadotropin subunit expression and salmon gonadotropin-releasing hormone (sgnrh) expression in the brain by quantitative real-time PCR. RNAseq revealed the expression of 75 known and 27 unknown salmonid olfactory genes of which 13 genes were differentially expressed between fish from the pre-spawning area and from the coastal area, suggesting an important role of these genes in homing. A clear progression towards final maturation was characterised by higher plasma 17α,20β-dihydroxy-4-pregnen-3-one (DHP) levels, increased pituitary luteinizing hormone β subunit (lhβ) expression and sgnrh expression in the post brain, and lower plasma testosterone (T) and 17β-estradiol (E2) levels. Olfactomedins and ependymin are candidates among the differentially expressed genes that may connect olfactory reception to the expression of sgnrh to regulate final maturation.     

Influence of aperiodic summer droughts on leaf litter breakdown and macroinvertebrate assemblages: testing the <Emphasis Type="Italic">drying memory</Emphasis> in a Central Apennines River (Aterno River,Italy)

Management of multiple exploited stocks of anadromous salmonids in large catchments requires understanding of movement and catchment use by the migrating fish and of their harvesting. The spawning migration of sea trout (Salmo trutta) and Atlantic salmon (Salmo salar) was studied in the River Tweed, UK, using acoustic telemetry to complement exploitation rate data and to quantify catchment penetration. Salmon (n = 79) and sea trout (n = 65) were tagged in the tidal-influenced Tweed in summer–autumn. No tagged salmon left the river before spawning, but 3% (2010) and 8% (2011) of pre-spawning sea trout dropped out. Combined tag regurgitation/fish mortality in salmon was 12.5%, while trout mortality was 6% (2010) and 0% (2011). The estimated spawning positions of salmon and sea trout differed; tagged salmon were mostly in the main channel while trout occurred mostly in the upper Tweed and tributaries. Early fish migrated upstream slower than later fish, but sea trout moved through the lower-middle river more quickly than salmon, partly supporting the hypothesis that the lower exploitation rate in autumn of trout (1 vs 3.3% for salmon) there is generated by differences in migration behaviour.     

A test of the hypothesis of pheromone attraction in salmonid migration

Synopsis We tested the hypothesis that anadromous salmonids are guided on their homeward migration by population-specific pheromones. Our findings do not support the hypothesis. Wild migrant Arctic charr,Salvelinus alpinus, from Ikarut River, Labrador were transferred and held in a tributary previously uninhabited by anadromous fish. None of the charr migrating up Ikarut River entered the tributary after fish were transferred. Similarly, migrant charr, which were caught in Ikarut River and released in the tributary below the captive fish, did not remain in the tributary. We re-evaluated the data which have been used to uphold the concept of pheromone attraction in salmonid migration and concluded that support for the hypothesis is unsubstantiated.     

A review of the likely effects of climate change on anadromous Atlantic salmon Salmo salar and brown trout Salmo trutta,with particular reference to water temperature and flow

The present paper reviews the effects of water temperature and flow on migrations, embryonic development, hatching, emergence, growth and life‐history traits in light of the ongoing climate change with emphasis on anadromous Atlantic salmon Salmo salar and brown trout Salmo trutta. The expected climate change in the Atlantic is for milder and wetter winters, with more precipitation falling as rain and less as snow, decrease in ice‐covered periods and frequent periods with extreme weather. Overall, thermal limits for salmonids are species specific. Scope for activity and growth and optimal temperature for growth increase with temperature to an optimal point before constrain by the oxygen content of the water. The optimal temperature for growth decreases with increasing fish size and varies little among populations within species, whereas the growth efficiency may be locally adapted to the temperature conditions of the home stream during the growth season. Indirectly, temperature influences age and size at smolting through its effect on growth. Time of spawning, egg hatching and emergence of the larvae vary with temperature and selective effects on time of first feeding. Traits such as age at first maturity, longevity and fecundity decrease with increasing temperature whilst egg size increases with temperature. Water flow influences the accessibility of rivers for returning adults and speed of both upstream and downstream migration. Extremes in water flow and temperature can decrease recruitment and survival. There is reason to expect a northward movement of the thermal niche of anadromous salmonids with decreased production and population extinction in the southern part of the distribution areas, migrations earlier in the season, later spawning, younger age at smolting and sexual maturity and increased disease susceptibility and mortality. Future research challenges are summarized at the end of the paper.     

Habitat coupling in a large lake system: delivery of an energy subsidy by an offshore planktivore to the nearshore zone of Lake Superior

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Do Insectivorous Birds use Volatile Organic Compounds from Plants as Olfactory Foraging Cues? Three Experimental Tests

Some insectivorous birds orient towards insect‐defoliated trees even when they do not see the foliar damage or the herbivores. There are, however, only a few studies that have examined the mechanisms behind this foraging behaviour. Previous studies suggest that birds can use olfactory foraging cues (e.g. volatile organic compounds (VOCs) emitted by defoliated plants), indirect visual cues or a combination of the two sensory cues. VOCs from insect‐defoliated plants are known to attract natural enemies of herbivores, and researchers have hypothesized that VOCs could also act as olfactory foraging cues for birds. We conducted three experiments across a range of spatial scales to test this hypothesis. In each experiment, birds were presented with olfactory cues and their behavioural responses or foraging outcomes were observed. In the first experiment, two different VOC blends, designed to simulate the volatile emissions of mountain birch (Betula pubescens ssp. czerepanovii) after defoliation by autumnal moth (Epirrita autumnata) larvae, were used in behavioural experiments in aviaries with pied flycatchers (Ficedula hypoleuca). The second experiment was a field‐based trial of bird foraging efficiency; the same VOC blends were applied to mountain birches, silver birches (B. pendula) and European white birches (B. pubescens) with plasticine larvae attached to the trees to serve as artificial prey for birds and provide a means to monitor predation rate. In the third experiment, the attractiveness of silver birch saplings defoliated by autumnal moth larvae versus intact controls was tested with great tits (Parus major) and blue tits (Cyanistes caeruleus) in an aviary. Birds did not orient towards either artificial or real trees with VOC supplements or towards herbivore‐damaged saplings when these saplings and undamaged alternatives were hidden from view. These findings do not support the hypothesis that olfactory foraging cues are necessary in the attraction of birds to herbivore‐damaged trees.     

Small larvae in large rivers: observations on downstream movement of European grayling Thymallus thymallus during early life stages

Behaviour of early life stages of the salmonid European grayling Thymallus thymallus was investigated by assessing the timing of larval downstream movement from spawning areas, the depth at which larvae moved and the distribution of juvenile fish during summer in two large connected river systems in Norway. Trapping of larvae moving downstream and electrofishing surveys revealed that T. thymallus larvae emerging from the spawning gravel moved downstream predominantly during the night, despite light levels sufficient for orientation in the high‐latitude study area. Larvae moved in the water mostly at the bottom layer close to the substratum, while drifting debris was caught in all layers of the water column. Few young‐of‐the‐year still resided close to the spawning areas in autumn, suggesting large‐scale movement (several km). Together, these observations show that there may be a deliberate, active component to downstream movement of T. thymallus during early life stages. This research signifies the importance of longitudinal connectivity for T. thymallus in Nordic large river systems. Human alterations of flow regimes and the construction of reservoirs for hydropower may not only affect the movement of adult fish, but may already interfere with active movement behaviour of fish during early life stages.     

Turbidity Hampers Mate Choice in a Pipefish

European coastal waters have in recent years become more turbid as algal growth has increased, probably due to eutrophication, global warming and changes in fish communities. Turbidity reduces visibility, and such changes may in turn affect animal behaviour as well as evolutionary processes that are dependent on visual stimuli. In this study we experimentally manipulated water visibility and olfactory cues to investigate mate choice using the sex role‐reversed broad‐nosed pipefish Syngnathus typhle as our study organism. We show that males spent significantly longer time assessing females when they had access to full visual cues, compared to when visibility was reduced. Presence or absence of olfactory cues from females did not affect mate choice, suggesting that the possible use of smell could not make up for a reduction in visibility. This implies that mate choice is environmentally dependent and that an increased turbidity may affect processes of sexual selection through an impaired possibility for visually based mate choice.     

Oviposition preferences in Grapholita molesta: the relative importance of visual and olfactory cues

The oriental fruit moth, Grapholita molesta (Busck) (Lepidoptera: Tortricidae), is a major pest of fruit trees worldwide. Females oviposit on the young leaves of peach [Prunus persica (L.) Batsch (Rosaceae)] shoots at twilight, when light intensity changes markedly. Previous researches have shown that the oviposition response of this moth is guided particularly by visual and olfactory cues from host plants. However, the relative importance of visual and olfactory cues in their oviposition preference is largely unknown. As a crepuscular moth, how do choices change as light intensity drops rapidly from day to night? In the present study, through two‐choice behavioural experiments, the oviposition responses of G. molesta to visual and olfactory cues (alone or in combination) from peach shoots were tested at four light intensities ranging from daylight to starlight. Grapholita molesta showed similar oviposition responses to plant cues under 1 000, 100, 1 and 0.01 mW m^?2 illumination. Olfactory cues from peach shoots attracted significantly more oviposition than the negative control, whereas visual cues alone did not. Furthermore, olfactory cues were more attractive than visual cues, and no significant interaction was observed between the responses to the two cues. Our findings indicate that G. molesta females mainly rely on olfactory cues to recognize oviposition sites, regardless of differences in light intensity. These results do not provide evidence for attraction to visual cues, but the possibility that the brightness of leaves might be used to guide oviposition is discussed.     

Elevation of gene expression for salmon gonadotropin‐releasing hormone in discrete brain loci of prespawning chum salmon during upstream migration

Our previous studies suggested that salmon gonadotropin‐releasing hormone (sGnRH) neurons regulate both final maturation and migratory behavior in homing salmonids. Activation of sGnRH neurons can occur during upstream migration. We therefore examined expression of genes encoding the precursors of sGnRH, sGnRH‐I, and sGnRH‐II, in discrete forebrain loci of prespawning chum salmon, Oncorhynchus keta. Fish were captured from 1997 through 1999 along their homing pathway: coastal areas, a midway of the river, 4 km downstream of the natal hatchery, and the hatchery. Amounts of sGnRH mRNAs in fresh frozen sections including the olfactory bulb (OB), terminal nerve (TN), ventral telencephalon (VT), nucleus preopticus parvocellularis anterioris (PPa), and nucleus preopticus magnocellularis (PM) were determined by quantitative real‐time polymerase chain reactions. The amounts of sGnRH‐II mRNA were higher than those of sGnRH‐I mRNA, while they showed similar changes during upstream migration. In the OB and TN, the amounts of sGnRH mRNAs elevated from the coast to the natal hatchery. In the VT and PPa, they elevated along with the progress of final maturation. Such elevation was also observed in the rostroventral, middle, and dorsocaudal parts of the PM. The amounts of gonadotropin IIβ and somatolactin mRNAs in the pituitary also increased consistently with the elevation of gene expression for sGnRH. These results, in combination with lines of previous evidence, indicate that sGnRH neurons are activated in almost all the forebrain loci during the last phases of spawning migration, resulting in coordination of final gonadal maturation and migratory behavior to the spawning ground. © 2005 Wiley Periodicals, Inc. J Neurobiol, 2005     

Reproductive energy expenditure and changes in body morphology for a population of Chinook salmon Oncorhynchus tshawytscha with a long distance migration

Energetic demands of a long freshwater migration, extended holding period, gamete development and spawning were evaluated for a population of stream‐type Chinook salmon Oncorhynchus tshawytscha. Female and male somatic mass decreased by 24 and 21%, respectively, during migration and by an additional 18 and 12% during holding. Between freshwater entry and death after spawning, females allocated 14% of initial somatic energy towards gonad development and 78% for metabolism (46, 25 and 7% during migration, holding and spawning, respectively). Males used only 2% of initial somatic energy for gonad development and 80% on metabolic costs, as well as an increase in snout length (41, 28 and 11% during migration, holding and spawning, respectively). Individually marked O. tshawytscha took between 27 and 53 days to migrate 920 km. Those with slower travel times through the dammed section of the migration corridor arrived at spawning grounds with less muscle energy than faster migrants. Although energy depletion did not appear to be the proximate cause of death in most pre‐spawn mortalities, average final post‐spawning somatic energy densities were low at 3·6 kJ g^?1 in females and 4·1 kJ g^?1 in males, consistent with the concept of a minimum energy threshold required to sustain life in semelparous salmonids.     

Modeling monkeys: A comparison of computer-generated and naturally occurring foraging patterns in two species of neotropical primates

We present a series of computer-generated foraging models (random movement, olfactory navigation, and spatial memory) designed to examine the manner in which sensory cues and cognitive skills might be used by rainforest monkeys to locate patchily distributed feeding sites. These simulations are compared with data collected in the Amazon Basin of northeastern Peru on the foraging patterns of two species of neotropical primates, the moustached tamarin monkey (Saguinus mystax) and the saddle-back tamarin monkey (Saguinus fuscicollis). The results indicate that, although tamarins may rely on olfactory cues to locate nearby feeding sites, their foraging patterns are better explained by an ability to maintain a detailed spatial map of the location and distribution of hundreds of feeding trees in their home range. There is evidence that such informationis retained for a period of at least several weeks and is used to minimize the distance traveled between widely scattered feeding sites. The use of computer simulations provides a powerful research tool for generating predictive models regarding the role of memory and sensory cues in animal foraging patterns.     

Spawning migrations in landlocked Atlantic salmon: time series modelling of river discharge and water temperature effects

River discharge and water temperature are frequently cited as controlling the upstream migration of adult salmonids to their spawning areas. The results of earlier studies on the effect of these environmental factors were examined. The statistical methods employed in some of these studies failed to consider the serial correlation often found in migration time series. To assess the effects of discharge and temperature on the migratory activity of the landlocked Atlantic salmon (ouananiche, Salmo salar ), 12 years of data on spawning migrations in the Mistassini R., Quebec, Canada, were analysed and the results obtained by ordinary least squares regression and time series regression were compared. In six of the 12 years, upstream migratory movement was related negatively to changes in river flow, suggesting that fish favour falling water phases for ascent. Water temperature appeared to have little effect on migratory movement. The mean body size of migrating fish decreased significantly throughout the summer; early migrants were on average 11.4% larger (mean fork length 522mm) than late migrants (469 mm). Larger, 3-lake-year salmon migrated 7.2 days earlier than 2-lake-year salmon. Because the residuals from ordinary regression exhibited strong autocorrelation, time series regression was more appropriate than ordinary regression for the analysis of migration time Series.     

RHEOTROPISM IN FISHES

(1) The fluid properties of air and water enable animals to orientate to flow and this behaviour in water is termed rheotaxis. Fish, however, have a wide range of responses to currents, extending beyond simple orientation, and the term rheotropism is therefore used as a ‘portmanteau’ word to describe all such reactions. (2) Fish detect currents directly by flow over the body surface or indirectly by other stimuli. Indirect responses are more common and occur in response to visual, tactile and inertial stimuli resulting from displacement of the fish by the current. Reactions to displacement of visual images are called optomotor reactions. The lateral line is not involved except in the detection of small localized jets of water. It has not been demonstrated that any fish can detect the current by electrical stimuli, although it is theoretically possible for some to do so. (3) In the basic form of rhotaxis the fish heads upstream and maintains station by stemming the current. Current detection thresholds fall within the range 0.4 to 10 cm/s for tactile stimuli but may be as low as 0.03 cm/s for visual stimuli. (4) Visual responses have been studied by simulating displacement by the current in optomotor apparatus. Fish respond to a rotating black-and-white-striped background by compensatory movements of the head and eyes - optokinetic nystagmus - or by the optomotor reaction, in which the fish swims with the background. (5) Fish show an orthokinesis in optomotor apparatus, their mean swimming speed increasing with the speed of rotation of the background. The precise form of the relationship varies between species and there is also considerable individual variation in performance. Fish accelerate and decelerate relative to the background, fixating on a particular stripe for short periods. (6) Factors limiting the appearance of the optomotor response are contrast, illuminance, acuity, critical flicker fusion frequency and spectral sensitivity. (7) Fish tolerate retinal image movements equivalent to those received when they are carried forwards by the current but not to those received when they are carried backwards. There are ganglion cells in the optic tectum which are sensitive to the direction of movement of targets across the visual field. In the goldfish there are significantly more units sensitive to movements in the temporo-nasal than in the opposite direction. (8) There are close parallels between the behaviour of fish in schools and in an optomotor apparatus. The optomotor response is apparently innate, occurring in newly hatched fry. (9) Physical and chemical factors can modify rheotaxis. Temperature and olfactory stimuli affect both the sign of the taxis and the kinetic component of the behaviour. (10) Thyroid hormones which are involved in the control of migration have been shown to affect the kinetic component of rheotaxis. (11) Fish show a number of hydrodynamic adaptations to life in currents. Morphological modifications are greatest in fish from torrential streams, which show extreme dorsoventral flattening and have specialized adhesive organs. Other fish select areas of low velocity or decrease their buoyancy with increasing current speed. (12) Rheotropic behaviour plays an important role in the distribution of fish within stream systems, in the maintenance of territory and station and in feeding behaviour. Territory, station and spawning sites in salmonids are all selected in relation to water velocity. (13) Water currents are thought to provide either a transport system or directional clues for fish on migration. The fish either does not respond to the current and is carried passively downstream, or it makes an orientated movement, swimming up- or downstream. (14) Eggs and larvae are known to drift passively downstream from their spawning grounds and some adult fish may also drift passively. In the sea both adult and juvenile fish use a form of modulated drift associated with vertical migration. Fish move up into midwater either by direct tidal selection or in relation to the diel cycle of illuminance. In fresh water the downstream migrations of salmonid fry, and smolts under some conditions, occur by modulated drift. (15) There is no evidence that fish migrating in the sea orientate to the current, but in fresh water the upstream migrations of diadromous fish are clearly orientated movements. (16) Water velocity is a major factor for salmonids migrating upstream. For fry it limits the occurrence of upstream migrations and for adults it can also prevent upstream movement. But migrations are often initiated by freshets, and changing water velocity is thought to be the most important factor associated with a freshet. (17) Both environmental and genetic factors affect the direction of migration in relation to the current. In some sockeye salmon fry direction is determined by temperature, but in others the overall direction of movement is genetically determined and environmental factors only modify the behaviour. (18) Rheotropic behaviour has a number of important practical applications in the capture of fish and in guiding them past dams and power stations. (19) The optomotor response plays a basic role in the capture of roundfish by trawls under conditions when the fish can see the gear. Many fish are caught because they become fatigued after a prolonged period of swimming at the same speed as the trawl. (20) Most success in guiding fish away from hazardous areas and bypassing them round dams has been achieved with mechanical barriers which depend on rheotropic reactions of the fish. (21) Louvre screens are very successful in deflecting juvenile salmonids migrating downstream past small dams but are impracticable at large dams. Instead, the turbine intakes are commonly sited at a considerable depth and fish are bypassed by mechanical screens either at the surface of the forebay or into the gatewells immediately upstream of the turbine intakes. (22) With upstream migrants the basic problem is to attract fish to the lower end of the fishways. An adequate ‘attraction velocity’ is an important feature of fishways, which must be sited so that the fish avoid the high velocity discharges from spillways and turbines.