The influence of flow velocity on the response of rheophilic fish to visual cues.

The strong association with visual cues exhibited by fish that prefer to inhabit flowing water (rheophilic species) may help reduce the energetic costs of maintaining position due to the provision of spatial points of reference. If this "Station Holding Hypothesis" is true, a positive relationship between the association with visual cues and flow velocity is expected. This hypothesis was tested experimentally by quantifying the response of common minnow (Phoxinus phoxinus) and brown trout (Salmo trutta) to visual cues under three flow velocities. In contradiction to the prediction, there was no evidence that the association with strong visual cues was positively related to flow velocity when fish were presented with vertical black stripes in an open channel flume, although interspecific variation in response was observed. The association with visual cues was relatively weak in trout, compared to minnow that spent 660% more time associated with the zone in which visual cues were present during the treatment, than the control when visual cues were absent. Trout tended to be more exploratory and made short visits to the area where visual cues were present, whereas minnow associated with the cues for longer. The strong association with visual cues independent of flow velocity exhibited by minnow and the weak association across all velocities by trout suggest that this behaviour is unlikely to reflect a strategy to minimise the energetic cost of maintaining position in flowing water. Minnow may have used the visual cues as a proxy indicator of physical structure that provides alternative benefits, such as refuge from predators. Trout may have employed alternative cues (e.g. mechanosensory) to seek more energetically favourable regions of the experimental area, reducing the importance of stationary visual stimuli.

Response of upstream migrating juvenile European eel (Anguilla anguilla) to electric fields: Application of the marginal gains concept to fish screening.

The decline in European eel (Anguilla anguilla) recruitment over the past half-century is partly due to river infrastructure that delays or blocks upstream migration to rearing habitat. Stimuli, such as electricity, can be used to modify the behaviour of downstream moving fish and guide them to preferred routes of passage at river infrastructure; but research on upstream migrating juvenile eel remains limited. The response of upstream migrating juvenile eel exposed to pulsed direct current (PDC) electric fields was investigated using a recirculatory flume. Eel were presented a choice of two routes upstream under either: (1) a treatment condition, in which the selection of one route resulted in exposure to High Electric Field (HEF) strength that was between 1.5-2 times stronger than the Low Electric Field (LEF) strength encountered in the alternative route; or (2) a control in which the electric field was absent in both routes. Under the treatment, five different mean HEF strengths (0.53, 0.77, 1.22, 2.17 and 3.74 Vcm-1) were tested at one of two frequencies (2 and 10 Hz). Route choice, distance downstream of the first set of electrodes at which an initial response was observed and avoidance behaviours (acceleration, retraction, switching and rejection) were compared among treatments. For the 1.22, 2.17 and 3.74 Vcm-1 and under 2 Hz, eel preferred to pass the LEF route. Avoidance was greater in the HEF route and positively related to field strength. The distance of the initial response did not differ between routes, field strengths or frequency. Upstream migrating eel avoided electric fields indicating potential to develop this approach for fish guidance. Further work is needed to test prototypes in field settings, particularly in combination with traditional physical screens to water intakes as part of a process of applying the concept of marginal gains to advance environmental impact mitigation technology.

The response of common carp (Cyprinus carpio) to insonified bubble curtains.

Acoustic bubble curtains have been marketed as relatively low cost and easily maintained behavioural deterrents for fisheries management. Their energy efficiency can be improved by reducing air flow and exploiting bubble resonance. In a series of three flume experiments, we: (1) investigated the reactions of carp to a low air flow bubble curtain, (2) compared the effectiveness of resonant versus non-resonant insonified bubble curtains (for the same volume flux of gas injected through the nozzles) to deter passage, and determined the stimuli responsible for eliciting deterrence, and (3) included the effect of visual cues generated by the bubble curtain. This study showed that bubble curtains with a higher proportion of resonant bubbles deterred carp relatively better. Passage rejection was likely influenced by multiple cues at distances within a body length of the fish- specifically the rate of change in both particle motion and flow velocity caused by rising bubbles. All acoustic bubble curtains were less effective in the presence of daylight, suggesting that vision plays an important role at mediating carp reactions. We discuss the importance of ascertaining the bubble size distribution, in addition to the gas flow rate and aperture size, when characterising acoustically active bubble curtains.

Collective behaviour of the European minnow (Phoxinus phoxinus) is influenced by signals of differing acoustic complexity.

Collective behaviour, such as shoaling in fish, benefits individuals through a variety of activities such as social information exchange and anti-predator defence. Human driven disturbance (e.g. anthropogenic noise) is known to affect the behaviour and physiology of individual animals, but the disruption of social aggregations of fish remains poorly understood. Anthropogenic noise originates from a variety of activities and differs in acoustic structure, dominant frequencies, and spectral complexity. The response of groups of fish may differ greatly, depending on the type of noise, and how it is perceived (e.g. threatening or attractive). In a controlled laboratory study, high resolution video tracking in combination with fine scale acoustic mapping was used to investigate the response of groups of European minnows (Phoxinus phoxinus) to signals of differing acoustic complexity (sinewave tones vs octave band noise) under low (150 Hz) and high (2200 Hz) frequencies. Fish startled and decreased their mean group swimming speed under all four treatments, with low frequency sinewave tones having the greatest influence on group behaviour. The shoals exhibited spatial avoidance during both low frequency treatments, with more time spent in areas of lower acoustic intensity than expected. This study illustrates how noise can influence the spatial distribution and social dynamics within groups of fish, and owing to the high potential for freshwater aquatic environments to be influenced by anthropogenic activity, wider consequences for populations should be further investigated.

Artificial light at night (ALAN) affects the downstream movement behaviour of the critically endangered European eel, Anguilla anguilla.

Artificial light at night (ALAN) is considered one of the most pervasive forms of environmental pollution. It is an emerging threat to freshwater biodiversity and can influence ecologically important behaviours of fish. The European eel (Anguilla anguilla) is a critically endangered catadromous species that migrates downstream to the ocean to spawn in the Sargasso Sea. Given the pervasive nature of ALAN, many eel will navigate through artificially lit routes during their seaward migration, and although considered negatively phototactic, their response has yet to be quantified. We investigated the response of downstream moving European eel to simulated ALAN using a Light Emitting Diode unit in an experimental flume. We presented two routes of passage under: (1) a dark control (both channels unlit), (2) low ALAN (treatment channel lit to ca. 5 lx), or (3) high ALAN (treatment channel lit to ca. 20 lx). Eel were: (i) more likely to reject an illuminated route when exposed to high levels of ALAN; (ii) less likely to select the illuminated channel when given a choice; and (iii) passed downstream more rapidly when the illuminated route was selected. This study quantified the response of the critically endangered European eel to ALAN under an experimental setting, providing the foundations for future field based research to validate these findings, and offering insight on the ecological impacts of this major environmental pollutant and driver of global change.

A mechanical approach to understanding the impact of the nematode Anguillicoloides crassus on the European eel swimbladder.

One of the most detrimental factors in the drastic decline of the critically endangered European eel (Anguilla anguilla) was the inadvertent introduction of the invasive nematode Anguillicoloides crassus Infection primarily affects the swimbladder, a gas-filled organ that enables the eel to control its depth in the water. A reduction in swimbladder function may be fatal for eel undergoing their spawning migration to the Sargasso Sea, a journey of over 5000 km. Although the physiological damage caused by this invasive parasite is well studied through the use of quantifiable gross pathological indices, providing a good measure of the swimbladder health status, they cannot separate the role of mechanical and morphological damage. Our study examined the appropriateness of three commonly used indices as a measure of mechanical damage by performing uniaxial tensile tests on swimbladder specimens obtained from an infected eel population. When the test results were compared with the gross pathological indices it was found that thickness correlated most strongly with mechanical damage, both confirming and, more importantly, explaining the counterintuitive findings of earlier work. In a damaged swimbladder, the immune response leads to a trade-off; increasing wall thickness raises the pressure required for organ rupture but decreases strength. The results indicate that for moderate infection the mechanical integrity of the swimbladder can be maintained. For severe infection, however, a reduction in mechanical integrity may reach a tipping point, thereby affecting the successful completion of their oceanic migration.

COVID-19 provides an opportunity to advance a sustainable UK fisheries policy in a post-Brexit brave new world.

Brexit creates a systemic shock that provides a unique opportunity for the UK to implement a new sustainable Fisheries Policy to better manage the multiple stocks on which future fishers will depend on leaving the European Union. At the same time, the global slowdown of commercial fishing as a result of COVID-19 has reduced pressure on some threatened stocks to levels not seen since the Second World War. In combination, Brexit and the COVID-19 slowdown have created a unique opportunity to facilitate the recovery of a threatened resource. Nevertheless, challenges remain as fisheries represent only 0.12% of UK economic output, presenting a risk that opportunities for more sustainable management will be lost during wider trade negotiations. Reduced fishing pressure during the COVID-19 era will enable stocks an opportunity to recover if supported by a new UK Fisheries Policy that focuses on: (a) re-establishing the role of Maximum Sustainable Yield to set limits that enable the recovery of fish populations initiated during the COVID-19 era; (b) ensuring that catch targets are set with the aim to maintain biomass at 120% of that which will achieve Maximum Sustainable Yield; (c) improving coherent resource management that also considers the expensive use of carbon associated with unsustainable fishing, and the need to protect fish throughout their life-cycle; and (d) constructing and effectively enforcing protection of a resilient network of Marine Protected Areas despite potential protests from EU member states.

Group behavior and tolerance of Eurasian minnow (Phoxinus phoxinus) in response to tones of differing pulse repetition rate.

Behavioral guidance systems are commonly used in freshwater fish conservation. The biological relevance of sound to fish and recorded responses to human-generated noise supports the viability of the use of acoustics as an effective stimulus in such technologies. Relatively little information exists on the long-term responses and recovery of fish to repeated acoustic exposures. In a controlled laboratory study, the response and tolerance of Eurasian minnow (Phoxinus phoxinus) shoals to tonal signals (150 Hz of 1 s pulse duration) differing only in temporal characteristics ("continuous," "slow," "intermediate," or "fast" pulse repetition rate) were investigated. In comparison to independent control groups, fish increased their mean group swimming speed, decreased inter-individual distance, and became more aligned in response to the onset of all four acoustic treatments. The magnitude of response, and time taken to develop a tolerance to a treatment differed according to pulse repetition rate. Groups were found to have the greatest and longest lasting response to tone sequences tested in this study when they were pulsed at an intermediate rate of 0.2 s-1. This study illustrates the importance of understanding the response of fish to acoustic signals, and will assist toward the development of longer-term effective acoustic guidance systems.

Masking a fish's detection of environmental stimuli: application to improving downstream migration at river infrastructure.

According to Signal Detection Theory, the ability to detect a stimulus (discriminability, d') is inversely related to the magnitude of internal and external noise. In this study, downstream moving brown trout Salmo trutta were used to investigate whether external hydrodynamic noise (in this case turbulence) could mask a signal that induced an unwanted response, such as rejecting accelerating velocity gradients commonly encountered at entrances to fish bypass channels. S. trutta behaviour was quantified in the absence (control) or presence of an accelerating velocity gradient created by an unconstricted or constricted channel, respectively, under two levels (low and high) of background turbulent kinetic energy (hydrodynamic noise). Experiments were conducted in an indoor recirculating flume in the dark and a range of passage metrics were quantified. Under the control condition, most (ca. 91%) S. trutta passed, usually oriented downstream (67%), with minimal delay (median 0.13 min). In comparison, fewer S. trutta (ca. 43%) passed under constricted conditions, they tended to orient facing into the flow (ca. 64%) and delay was greater (median > 20 min). When viewed from a coarse-scale perspective, discriminability of the velocity gradient was lower when turbulence was high suggesting masking of the signal occurred. However, the resulting increase in the percentage of fish that passed, decrease in time to pass and reduction in the distance at which S. trutta reacted (switched orientation) was subtle and non-significant. Despite the mixed results obtained, the use of masking to manipulate an animal's perception of environmental stimuli as a fisheries management tool is conceptually valid and the results of this experiment present a useful stepping stone for future research.

Assessing hydrodynamic space use of brown trout, Salmo trutta, in a complex flow environment: a return to first principles.

It is commonly assumed that stream-dwelling fish should select positions where they can reduce energetic costs relative to benefits gained and enhance fitness. However, the selection of appropriate hydrodynamic metrics that predict space use is the subject of recent debate and a cause of controversy. This is for three reasons: (1) flow characteristics are often oversimplified, (2) confounding variables are not always controlled and (3) there is limited understanding of the explanatory mechanisms that underpin the biophysical interactions between fish and their hydrodynamic environment. This study investigated the space use of brown trout, Salmo trutta, in a complex hydrodynamic flow field created using an array of different sized vertically oriented cylinders in a large open-channel flume in which confounding variables were controlled. A hydrodynamic drag function (D) based on single-point time-averaged velocity statistics that incorporates the influence of turbulent fluctuations was used to infer the energetic cost of steady swimming. Novel hydrodynamic preference curves were developed and used to assess the appropriateness of D as a descriptor of space use compared with other commonly used metrics. Zones in which performance-enhancing swimming behaviours (e.g. Kármán gaiting, entraining and bow riding) that enable fish to hold position while reducing energetic costs (termed 'specialised behaviours') were identified and occupancy was recorded. We demonstrate that energy conservation strategies play a key role in space use in an energetically taxing environment with the majority of trout groups choosing to frequently occupy areas in which specialised behaviours may be adopted or by selecting low-drag regions.

Response of seaward-migrating European eel (Anguilla anguilla) to manipulated flow fields.

Anthropogenic structures (e.g. weirs and dams) fragment river networks and restrict the movement of migratory fish. Poor understanding of behavioural response to hydrodynamic cues at structures currently limits the development of effective barrier mitigation measures. This study aimed to assess the effect of flow constriction and associated flow patterns on eel behaviour during downstream migration. In a field experiment, we tracked the movements of 40 tagged adult European eels (Anguilla anguilla) through the forebay of a redundant hydropower intake under two manipulated hydrodynamic treatments. Interrogation of fish trajectories in relation to measured and modeled water velocities provided new insights into behaviour, fundamental for developing passage technologies for this endangered species. Eels rarely followed direct routes through the site. Initially, fish aligned with streamlines near the channel banks and approached the intake semi-passively. A switch to more energetically costly avoidance behaviours occurred on encountering constricted flow, prior to physical contact with structures. Under high water velocity gradients, fish then tended to escape rapidly back upstream, whereas exploratory 'search' behaviour was common when acceleration was low. This study highlights the importance of hydrodynamics in informing eel behaviour. This offers potential to develop behavioural guidance, improve fish passage solutions and enhance traditional physical screening.

Managing sustainable development conflicts: the impact of stakeholders in small-scale hydropower schemes.

The growing importance of the environment and its management has simultaneously emphasized the benefits of hydroelectric power and its environmental costs. In a changing policy climate, giving importance to renewable energy development and environmental protection, conflict potential between stakeholders is considerable. Navigation of conflict determines the scheme constructed, making sustainable hydropower a function of human choice. To meet the needs of practitioners, greater understanding of stakeholder conflict is needed. This paper presents an approach to illustrate the challenges that face small-scale hydropower development as perceived by the stakeholders involved, and how they influence decision-making. Using Gordleton Mill, Hampshire (UK), as an illustrative case, soft systems methodology, a systems modeling approach, was adopted. Through individual interviews, a range of problems were identified and conceptually modeled. Stakeholder bias towards favoring economic appraisal over intangible social and environmental aspects was identified; costs appeared more influential than profit. Conceptual evaluation of the requirements to meet a stakeholder-approved solution suggested a complex linear systems approach, considerably different from the real-life situation. The stakeholders introduced bias to problem definition by transferring self-perceived issues onto the project owner. Application of soft systems methodology caused a shift in project goals away from further investigation towards consideration of project suitability. The challenge of sustainable hydropower is global, with a need to balance environmental, economic, and social concerns. It is clear that in this type of conflict, an individual can significantly influence outcomes; highlighting the need for more structured approaches to deal with stakeholder conflicts in sustainable hydropower development.