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A warming climate poses a fundamental problem for embryos that develop within eggs because their demand for oxygen (O 2 ) increases much more rapidly with temperature than their capacity for supply, which is constrained by diffusion across the egg surface. Thus, as temperatures rise, eggs may experience O 2 limitation due to an imbalance between O 2 supply and demand. Here, we formulate a mathematical model of O 2 limitation and experimentally test whether this mechanism underlies the upper thermal tolerance in large aquatic eggs. Using Chinook salmon ( Oncorhynchus tshawytscha ) as a model system, we show that the thermal tolerance of eggs varies systematically with features of the organism and environment. Importantly, this variation can be precisely predicted by the degree to which these features shift the balance between O 2 supply and demand. Equipped with this mechanistic understanding, we predict and experimentally confirm that the thermal tolerance of these embryos in their natural habitat is substantially lower than expected from laboratory experiments performed under normoxia. More broadly, our biophysical model of O 2 limitation provides a mechanistic explanation for the elevated thermal sensitivity of fish embryos relative to other life stages, global patterns in egg size and the extreme fecundity of large teleosts.

Author(s): Lehman, Brendan M.; Johnson, Rachel C.; Adkison, Mark; Burgess, Oliver T.; Connon, Richard E.; Fangue, Nann A.; Foott, J. Scott; Hallett, Sascha L.; Martinez–Lopez, Beatriz; Miller, Kristina M.; Purcell, Maureen K.; Som, Nicholas A.; Valdes–Donoso, Pablo; Collins, Alison L. | Abstract: Chinook Salmon (Oncorhynchus tshawytscha) are increasingly vulnerable to anthropogenic activities and climate change, especially at their most southern species range in California’s Central Valley. There is considerable interest in understanding stressors that contribute to population decline and in identifying management actions that reduce the impacts of those stressors. Along the west coast of North America, disease has been linked to declining numbers of salmonids and identified as a key stressor resulting in mortality. In the Central Valley, targeted studies have revealed extremely high prevalence of infectious agents and disease. However, there has been insufficient monitoring to understand the effect that disease may have on salmon populations. In order to inform future research, monitoring, and management efforts, a two-day workshop on salmon disease was held at UC Davis on March 14-15, 2018. This paper summarizes the science presented at this workshop, including the current state of knowledge of salmonid disease in the Central Valley and current and emerging tools to better understand its impacts on salmon. We highlight case studies from other systems where successful monitoring programs have been implemented. First, in the Klamath River where the integration of several data collection and modeling approaches led to the development of successful management actions, and second in British Columbia where investment in researching novel technologies led to breakthroughs in the understanding of salmon disease dynamics. Finally, we identify key information and knowledge gaps necessary to guide research and management of disease in Central Valley salmon populations.

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Animal populations are frequently infected by pathogens, but it is not always easy to determine the importance of pathogens to overall population dynamics. It is especially difficult to detect the effects of disease in population time series data because the effects are often local while overall population dynamics are also affected by larger-scale environmental factors. We overcame this difficulty by applying multivariate time series analysis to extract local effects from spawning abundance data and by comparing the survival rate of juvenile fall-run Chinook salmon Oncorhynchus tshawytscha from two locations in the Klamath River basin of California, one of which is affected by a high concentration of the myxozoan parasite Ceratomyxa shasta. To assess the effect of the disease (ceratomyxosis) caused by C. shasta on the population dynamics, we analyzed spatially structured abundance data for naturally spawning salmon and survival data for hatchery-released salmon for associations with exposure to ...

The parasite Ceratomyxa shasta has been implicated as a significant source of salmonid mortality in the lower Klamath River, California (i.e., below Iron Gate dam). A study of the prevalence of C. shasta and its geographic and temporal distribution throughout the Klamath River basin was conducted to determine when and where juvenile salmonids encounter lethal parasite doses. Susceptible rainbow trout Oncorhynchus mykiss were exposed to C. shasta 3–4 d at seven locations in the Klamath River between Beaver Creek and Keno Reservoir in April, June, July, September, and November 2003. Individuals from a Klamath River strain of fall Chinook salmon O. tshawytscha were held in three locations in the upper Klamath River in April, June, and July. In June 2004, rainbow trout were exposed to the parasite for 4 d at 18 locations from Klamath Lake to the mouth of the Klamath River, including several major spawning tributaries; one exposure occurred in the lower Klamath River. Rainbow trout mortality due to in...

The Lost River sucker (Deltistes luxatus) is a federally listed, endangered species inhabiting the hypereutrophic waters of Upper Klamath Lake in southern Oregon, USA. High pH (or =10) and elevated ammonia concentrations (or =1 mg NH(3)-N/L) often occur during blooms of cyanobacteria (Aphanizomenon flos-aquae) in the lake, with major fish kills sometimes following a mid- or late-summer "crash" of the cyanobacterial population. Previous histopathology analyses and bacterial sampling indicated that infections of the pathogenic bacterium Flavobacterium columnare might contribute to the fish kills. We hypothesized that prior exposure to adverse water quality conditions increases the susceptibility of Lost River suckers to F. columnare infections. To test this, we exposed juvenile Lost River suckers to four sublethal ammonia concentrations at pH 9.4 for 62 d. On day 31, fish in half of the aquaria were exposed to F. columnare. As expected, survival of the Lost River suckers decreased in aquaria inoculated with F. columnare. Ninety-four percent of the fish that died were infected by F. columnare in the gills, kidney, or skin, whereas none of the survivors or unexposed control fish was infected. However, contrary to our hypothesis, survival of the fish exposed to F. columnare increased significantly (p0.05) as unionized ammonia concentrations increased. Our results suggest that complex interactions can complicate prediction of the responses of fish to concurrent chemical stressors and bacterial pathogens.

One-year steelhead trout were reared at Coleman National Fish Hatchery, CA, USA. During the final 5 months of rearing, fish were maintained in either traditional concrete raceways or moved to a pond. Rearing conditions in the pond included a cobble bottom, structures, and surface disruption. In August, prior to splitting juveniles into raceway and pond groups, length, weight, gill Na + ,K + -ATPase activity, seawater tolerance and plasma levels of growth hormone (GH) and insulin-like growth factor I (IGF-I) were measured. On November 30th and January 3rd (prior to release on January 8th), fish were sampled as in August. The pond-reared fish grew faster (0.51 g day −1 from August to November and 0.66 g day −1 from November to January) than raceway-reared fish (0.41 and 0.26 g day −1 ) fed at similar rations (2% body weight day −1 ) through November and when raceway fish were fed more in December. In both November and January, pond-reared fish had significantly higher plasma GH levels and tolerated seawater transfer better than fish reared in raceways. Factors such as feeding conditions, structure, and fish density in the pond that differed from raceway conditions may have provided naturalization of the hatchery environment resulting in more physiologically competent smolts.

This study examined the hypothesis that exposure to sublethal concentrations of contaminants alters the gonadal condition of feral common carp Cyprinus carpio. Adult common carp in Lake Mead, Nevada, were collected from a contaminated site (Las Vegas Bay) that receives municipal and industrial effluent and from a reference site (Overton Arm) with a relatively low level of contamination. Fish were sampled seven times over a 1-year period extending over two separate spawning seasons. Morphometric and histopathological parameters of gonadal and germ cell development were determined. In males, the pattern of seasonal changes in the gonadosomatic index (GSI) was similar between the sites and showed no clear association with site-specific seasonal temperature profiles. However, Las Vegas Bay males had consistently lower GSI values and, on one of the sampling dates, a lower proportion of sperm relative to other germ cell stages (determined histologically). Further, Las Vegas Bay males had a higher incid...

With the increasing appreciation of global warming impacts on ecological systems, in addition to the myriad of land management effects on water quality, the number of literature citations dealing with the effects of water temperature on freshwater fish has escalated in the past decade. Given the many biological scales at which water temperature effects have been studied, and the growing need to integrate knowledge from multiple disciplines of thermal biology to fully protect beneficial uses, we held that a survey of the most promising recent developments and an expression of some of the remaining unanswered questions with significant management implications would best be approached collectively by a diverse research community. We have identified five specific topic areas of renewed research where new techniques and critical thought could benefit coldwater stream fishes (particularly salmonids): molecular, organism, population/species, community and ecosystem, and policy issues in water quality. Our hope is that information gained through examination of recent research fronts linking knowledge at various scales will prove useful in managing water quality at a basin level to protect fish populations and whole ecosystems. Standards of the past were based largely on incipient lethal and optimum growth rate temperatures for fish species, while future standards should consider all integrated thermal impacts to the organism and ecosystem.

The distribution and abundance of the myxosporean parasite Parvicapsula minibicornis in the Klamath River mirrored that of Ceratomyxa shasta, with which it shares both its vertebrate and invertebrate host. Assay of fish held at sentinel sites and water samples collected from those sites showed that parasite prevalence was highest below Iron Gate dam, which is the barrier to anadro- mous salmon passage. Above this barrier parasite levels fluctuated, with the parasite detected in the free-flowing river reaches between reservoirs. This was consistent with infection prevalence in the polychaete host, Manayunkia speciosa, which was greater than 1% only in populations tested below Iron Gate dam. Although a low prevalence of infection was detected in juvenile out-migrant fish in the Trinity River, the tributaries tested did not appear to be a significant source of the parasite to the mainstem despite the presence of large numbers of infected adult salmon that migrate and spawn there. Rainbow trout became infected during sentinel exposure, which expands the host range for P. minibicornis and suggests that wild rainbow trout populations are a reservoir for infection, especially above Iron Gate dam. High parasite prevalence in the lower Klamath River is likely a combined effect of high spore input from heavily infected, spawned adult salmon and the proximity to dense popula- tions of polychaetes.

Infectious Ceratomyxa shasta and Parvicapsula minibicornis actinospores were present in Klamath River samples collected in April, May, and June 2005. Juvenile Chinook salmon Oncorhynchus tshawytscha exposed to river water maintained at the ambient Klamath River temperature for 0, 4, 24, 72, and 168 h (7 d) developed asymptomatic infections from both parasites. Elevated water temperature (18 degrees C) in June may have reduced actinospore viability, as both C. shasta and P. minibicornis infection markedly declined in fish exposed for over 72 h. As judged by the prevalence of infection for both parasites, the number of infectious actinospores tended to increase or remain steady through the spring. Ceratomyxa shasta infections were characterized by the presence of a few trophozoites within granulomatous foci in mesentery adipose tissue and were consistently observed outside of the intestine. Similarly, low numbers of P. minibicornis were observed in kidney glomeruli and tubules but were not associated with inflammation. Parvicapsula minibicornis DNA was consistently detected by quantitative real-time polymerase chain reaction in filtered water samples collected each month and from each time posttransfer. These data and the high prevalence of infection observed in the exposed fish indicate that P. minibicornis actinospores were at a relatively high concentration in the river during the spring. In contrast, C. shasta DNA was only detected in half of the water sample sets and its detection did not correspond well to C. shasta infectivity. An approximately threefold increase in river flow from the April to the May water collection was not associated with a decline in either the detection of actinospores (particularly for P. minibicornis) or the prevalence of infection for both parasites. Actinospores of these myxosporean parasites have the potential to infect salmonids for at least 7 d after release from the alternate polychaete host.

Blood parameters, disease resistance, and the immune response were sequentially evaluated in rainbow trout Oncorhynchus mykiss with proliferative kidney disease (PKD). The fish were maintained under laboratory conditions, and the study group went through a full cycle of the disease. Hematological and serological changes occurred primarily in those fish with severe kidney lesions. Fish infected with the parasite that causes PKD demonstrated a greater resistance to bacterial challenge, and their immune responses were heightened when compared with those of uninfected fish. These data suggest that PKD alone is not a predisposing factor for secondary infections if the fish does not incur severe renal lesions.

The dynamics of IHNV infection and disease were followed in a juvenile Chinook salmon population both during hatchery rearing and for two weeks post-release. Cumulative weekly mortality increased from 0.03%–3.5% as the prevalence of viral infection increased from 2%–22% over the same four-week period. The majority of the infected salmon was asymptomatic. Salmon demonstrating clinical signs of infection shed 1000 pfu mL-1 of virus into the water during a 1 min observation period and had a mean concentration of 106 pfu mL-1 in their mucus. The high virus concentration detected in mucus suggests that it could act as an avenue of transmission in high density situations where dominance behavior results in nipping. Infected smolts that had migrated 295 km down river were collected at least two weeks after their release. The majority of the virus positive smolts was asymptomatic. A series of transmission experiments was conducted using oral application of the virus to simulate nipping, brief low dose waterborne challenges, and cohabitation with different ratios of infected to naïve fish. These studies showed that asymptomatic infections will occur when a salmon is exposed for as little as 1 min to >102 pfu mL-1, yet progression to clinical disease is infrequent unless the challenge dose is >104 pfu mL-1. Asymptomatic infections were detected up to 39 d post-challenge. No virus was detected by tissue culture in natural Chinook juveniles cohabitated with experimentally IHNV-infected hatchery Chinook at ratios of 1:1, 1:10, and 1:20 for either 5 min or 24 h. Horizontal transmission of the Sacramento River strain of IHNV from infected juvenile hatchery fish to wild cohorts would appear to be a low ecological risk. The study results demonstrate key differences between IHNV infections as present in a hatchery and the natural environment. These differences should be considered during risk assessments of the impact of IHNV infections on wild salmon and trout populations.

The anadromous fish health monitoring program began in full operation in January 1988 after the hiring of the lead pathologist. This short operating period limits the amount of information available at the time of this writing. Pre-release sampling of smolts revealed the presence of several sub-clinical pathogens. Organosomatic analysis results demonstrated no major abnormalities in the examined stocks. The results of the 1988 steelhead broodstock sampling are still pending.

This report documents the progress in the assigned tasks which have occurred during the second year of the Augmented Anadromous Fish Health Monitoring Project. Fish at seven Idaho Department of Fish and Game facilities were monitored for various pathogens and organosomatic analyses were performed on smolts prior to their release in the Spring of 1989. A disease database has been developed and facility impediments to fish health have been identified.