Jane Behrens

Oceanic Boundary Conditions for Jakobshavn Glacier. Part I: Variability and Renewal of Ilulissat Icefjord Waters, 2001–14*

JakobshavnGlacier, west Greenland, has respondedtotemperaturechangesinIlulissatIcefjord,intowhich it terminates.ThisstudycollectedhydrographicobservationsinsideIlulissatIcefjordandfromadjacentDiskoBay between2001and2014.ThewarmestdeepDiskoBaywaterswereblockedbytheentrancesillanddidnotreach Jakobshavn Glacier. In the fjord basin, the summer mean temperature was 2.88C from 2009 to 2013, excluding 2010, whenitwas 18C cooler. Despitethisvariability,summerpotential densities in the basinwereinthe narrow range of 27.20 # su # 27.31kgm 23 , and basin water properties matched those of Disko Bay in this layer each summer. This relation has likely held since at least 1980. Basin waters from 2009 and 2011‐13 were therefore similar tothosein1998/99, whenJakobshavn Glacier began toretreat, whilebasin watersin2010wereascoolas in the 1980s. The 2010 basin temperature anomaly was advected into Disko Bay, not produced by local atmospheric variability. This anomaly also shows that Ilulissat Icefjord basin waters were renewed annually or faster. Time series fragments inside the fjord did not capture the 2010 anomaly but show that the basin temperatures varied little subannually, outside of summer. Fjord velocity profiles from summer 2013 implied a basin renewal time scale of about 1 month. In model simulations of the fjord circulation, subglacial discharge from Jakobshavn Glacier could drive renewal of the fjord basin over a single summer, while baroclinic forcing from outside the fjord could not, because of the sill at the mouth.

The effect of temperature and body size on metabolic scope of activity in juvenile Atlantic cod Gadus morhua L.

Changes in ambient temperature affect the physiology and metabolism and thus the distribution of fish. In this study we used intermittent flow respirometry to determine the effect of temperature (2, 5, 10, 15 and 20°C) and wet body mass (BM) (~30-460g) on standard metabolic rate (SMR, mgO2h(-1)), maximum metabolic rate (MMR, mgO2h(-1)) and metabolic scope (MS, mgO2h(-1)) of juvenile Atlantic cod. SMR increased with BM irrespectively of temperature, resulting in an average scaling exponent of 0.87 (0.82-0.92). Q10 values were 1.8-2.1 at temperatures between 5 and 15°C but higher (2.6-4.3) between 2 and 5°C and lower (1.6-1.4) between 15 and 20°C in 200 and 450g cod. MMR increased with temperature in the smallest cod (50g) but in the larger cod MMR plateaued between 10, 15 and 20°C. This resulted in a negative correlation between the optimal temperature for MS (Topt) and BM, Topt being respectively 14.5, 11.8 and 10.9°C in a 50, 200 and 450g cod. Irrespective of BM cold water temperatures resulted in a reduction (30-35%) of MS whereas the reduction of MS at warm temperatures was only evident for larger fish (200 and 450g), caused by plateauing of MMR at 10°C and above. Warm temperatures thus seem favourable for smaller (50g) juvenile cod, but not for larger conspecifics (200 and 450g).

Effects of Hypoxic Exposure during Feeding on SDA and Postprandial Cardiovascular Physiology in the Atlantic Cod, Gadus morhua

Some Atlantic cod in the Bornholm Basin undertake vertical foraging migrations into severely hypoxic bottom water. Hypoxic conditions can reduce the postprandial increase in gastrointestinal blood flow (GBF). This could subsequently postpone or reduce the postprandial increase in oxygen consumption (MO2), i.e. the SDA, leading to a disturbed digestion. Additionally, a restricted oxygen uptake could result in an oxygen debt that needs to be compensated for upon return to normoxic waters and this may also affect the ability to process the food. Long-term cardio-respiratory measurements were made on fed G. morhua in order to understand how the cardio-respiratory system of feeding fish respond to a period of hypoxia and a subsequent return to normoxia. These were exposed to 35% water oxygen saturation for 90 minutes, equivalent to the time and oxygen level cod voluntarily endure when searching for food in the Bornholm Basin. We found that i) gastric and intestinal blood flows, cardiac output and MO2 increased after feeding, ii) gastric and intestinal blood flows were spared in hypoxia, and iii) there were no indications of an oxygen debt at the end of the hypoxic period. The magnitude and time course of the measured variables are similar to values obtained from fish not exposed to the hypoxic period. In conclusion, when cod in the field search for and ingest prey under moderate hypoxic conditions they appear to stay within safe limits of oxygen availability as we saw no indications of an oxygen debt, or negative influence on digestive capacity, when simulating field observations.

Evaluating dispersal potential of an invasive fish by the use of aerobic scope and osmoregulation capacity

Non-indigenous species (NIS) can impact marine biodiversity and ecosystem structure and function. Once introduced into a new region, secondary dispersal is limited by the physiology of the organism in relation to the ambient environment and by complex interactions between a suite of ecological factors such as presence of predators, competitors, and parasites. Early prediction of dispersal potential and future ‘area of impact’ is challenging, but also a great asset in taking appropriate management actions. Aerobic scope (AS) in fish has been linked to various fitness-related parameters, and may be valuable in determining dispersal potential of aquatic invasive species in novel environments. Round goby, Neogobius melanostomus, one of the most wide-ranging invasive fish species in Europe and North America, currently thrives in brackish and fresh water, but its ability to survive in high salinity waters is unknown to date. We show that AS in round goby is reduced by 30% and blood plasma osmolality increased (indicating reduced capacity for osmoregulation) at salinities approaching oceanic conditions, following slow ramping (5 PSU per week) and subsequent long-term acclimation to salinities ranging between 0 and 30 PSU (8 days at final treatment salinities before blood plasma osmolality measurements, 12–20 additional days before respirometry). Survival was also reduced at the highest salinities yet a significant proportion (61%) of the fish survived at 30 PSU. Reduced physiological performance at the highest salinities may affect growth and competitive ability under oceanic conditions, but to what extent reduced AS and osmoregulatory capacity will slow the current 30 km year-1 rate of advance of the species through the steep salinity gradient from the brackish Baltic Sea and into the oceanic North Sea remains speculative. An unintended natural experiment is in progress to test whether the rate of advance slows down. At the current rate of advance the species will reach the oceanic North Sea by 2018/2019, therefore time for taking preventative action is short.

Haemoglobin genotypes in cod (Gadus morhua L): Their geographic distribution and physiological significance

Haemoglobin polymorphism in cod (Gadus morhua L) has been investigated throughout the last 50years. Field studies have shed light on the geographic distribution of the two common alleles (HbI(1) and HbI(2)), and laboratory studies have shown effects of genotype on physiological traits such as growth, reproduction and hypoxia tolerance. The geographic distribution of alleles shows a correlation with temperature, with increasing frequency of HbI(1) in warmer areas. This is likely due to temperature-related differences in oxygen affinity of the three genotypes. We provide a general ecological introduction to cod haemoglobin polymorphism and a detailed discussion of physiological studies, particularly laboratory growth studies. Although differences in oxygen uptake are almost certainly a contributory mechanism to observed differences in traits such as growth rate, many other environmental, behavioural and social factors may also contribute, making it difficult to quantify the effect of HbI either experimentally or in the field.

The parasitic copepod Lernaeocera branchialis negatively affects cardiorespiratory function in Gadus morhua

The parasitic copepod Lernaeocera branchialis had a negative effect on cardiorespiratory function in Atlantic cod Gadus morhua such that it caused pronounced cardiac dysfunction with irregular rhythm and reduced stroke amplitude compared with uninfected fish. In addition, parasite infection depressed the postprandial cardiac output and oxygen consumption.

Consequences of eye fluke infection on anti-predator behaviours in invasive round gobies in Kalmar Sound

Larvae of the eye fluke, Diplostomum, emerge from snails and infect fish by penetrating skin or gills, then move to the lens where they may impair the vision of the fish. For the fluke to reproduce, a bird must eat the infected fish, and it has been suggested that they therefore actively manipulate the fish’s behaviour to increase the risk of predation. We found that round gobies Neogobius melanostomus, a species that was recently introduced to the Kalmar Sound of the Baltic Sea, had an eye fluke prevalence of 90–100%. We investigated how the infection related to behavioural variation in round gobies. Our results showed that the more intense the parasite-induced cataract, the weaker the host’s response was to simulated avian attack. The eye flukes did not impair other potentially important anti-predator behaviours, such as shelter use, boldness and the preference for shade. Our results are in accordance with the suggestion that parasites induce changes in host behaviour that will facilitate transfer to their final host.

Recovery of gastric evacuation rate in Atlantic cod Gadus morhua L surgically implanted with a dummy telemetry device

The current study investigated how the gastric evacuation rate (GER) was affected after surgically introducing dummies of a blood flow biotelemetry system into the abdominal cavity of Atlantic cod, Gadus morhua. Gastric evacuation experiments were performed two and 10 days postsurgery on surgically implanted and control G. morhua force-fed sandeel, Ammodytes tobianus. The results were compared with previously obtained estimates from unstressed conspecifics voluntarily feeding on a similar diet. After two days, GER was significantly lower in the group of fish with the dummy implants compared with the control group, but following 10 days of recovery no significant difference was seen between the two groups. The difference between implanted and control fish observed two days postsurgery may have resulted either from surgery, postsurgical stress and/or the presence of the implant. The conclusion is that 10 days of postsurgical recovery will stabilize GER in G. morhua, thus indicating that at this point the implant per se did not affect GER. Both the fish with surgical implants and controls in this study evacuated their stomachs much slower and with much higher interindividual variation compared with G. morhua feeding voluntarily on similar prey items. The lower GER and higher interindividual variation for force-fed fish indicate that handling, anaesthetization and force-feeding impair GER and that individual fish respond differently to the suppressing effects.

Early engagement of stakeholders with individual-based modelling can inform research for improving invasive species management : the round goby as a case study

Individual-based models (IBMs) incorporating realistic representations of key range-front processes such as dispersal can be used as tools to investigate the dynamics of invasive species. Managers can apply insights from these models to take effective action to prevent further spread and prioritize measures preventing establishment of invasive species. We highlight here how early-stage IBMs (constructed under constraints of time and data availability) can also play an important role in defining key research priorities for providing key information on the biology of an invasive species in order that subsequent models can provide robust insight into potential management interventions. The round goby, Neogobius melanostomus, is currently spreading through the Baltic Sea, with major negative effects being reported in the wake of its invasion. Together with stakeholders, we parameterize an IBM to investigate the goby’s potential spread pattern throughout the Gulf of Gdansk and the Baltic Sea. Model parameters were assigned by integrating information obtained through stakeholder interaction, from scientific literature, or estimated using an inverse modelling approach when not available. IBMs can provide valuable direction to research on invasive species even when there is limited data and/or time available to parameterize/fit them to the degree to which we might aspire in an ideal world. Co-development of models with stakeholders can be used to recognize important invasion patterns, in addition to identifying and estimating unknown environmental parameters, thereby guiding the direction of future research. Well-parameterized and validated models are not required in the earlier stages of the modelling cycle where their main utility is as a tool for thought.