Id |
Subject |
Object |
Predicate |
Lexical cue |
T1 |
0-85 |
Epistemic_statement |
denotes |
Fish disease dynamics in changing rivers: Salmonid Ceratomyxosis in the Klamath River |
T2 |
96-236 |
Epistemic_statement |
denotes |
The myxozoan parasite Ceratomyxa shasta has been identified as the main contributor of mortality in salmon in the Klamath River, California. |
T3 |
237-323 |
Epistemic_statement |
denotes |
The life cycle of the parasite is complex, involving a polychaete and a salmonid host. |
T4 |
742-967 |
Epistemic_statement |
denotes |
The analysis of the model shows that for current climate conditions, additional dam release can reduce the actinospore concentration up to 48% and the prevalence up to 40% thus providing a potential disease management option. |
T5 |
968-1148 |
Epistemic_statement |
denotes |
However, the infection risk is likely to increase for future climate conditions by 10-54% and this will lead to an infection level comparable to that of a recent high-disease year. |
T6 |
1292-1559 |
Epistemic_statement |
denotes |
We show that our detailed model system can be reduced to a simpler exponential dose-response function, which predicts infection levels based on transmission rates, travel time and mean spore concentration, quantities that can be measured in-situ or given empirically. |
T7 |
1560-1640 |
Epistemic_statement |
denotes |
The dose-response function may therefore be a useful tool in disease management. |
T8 |
1642-1852 |
Epistemic_statement |
denotes |
Myxozoans are a group of aquatic parasites that are known to cause several fish diseases that can have severe economic impact on fisheries and aquaculture (Hallett and Bartholomew, 2011; Okamura et al., 2015) . |
T9 |
2033-2140 |
Epistemic_statement |
denotes |
Several emerging diseases caused by myxozoans have been linked with climate change (Okamura et al., 2015) . |
T10 |
2141-2264 |
Epistemic_statement |
denotes |
The life cycle of myxozoans is complex and involves both invertebrate (mainly annelids) and vertebrate hosts (mainly fish). |
T11 |
2727-2873 |
Epistemic_statement |
denotes |
The parasite can be found throughout the Pacific Northwest, but its negative impact on native salmon populations is greatest in the Klamath River. |
T12 |
3190-3318 |
Epistemic_statement |
denotes |
However, quantifying the impact on wild fish populations is difficult due to the lack of long-term data (Okamura et al., 2015) . |
T13 |
4213-4470 |
Epistemic_statement |
denotes |
Future climate scenarios in the Klamath River predict significant differences in both temperature and precipitation patterns compared to current conditions (Perry et al., 2011; Ray et al., 2015) and are likely to affect spore concentration and disease risk. |
T14 |
4633-4777 |
Epistemic_statement |
denotes |
Therefore, determining the effect of changing environmental conditions on the disease dynamics is crucial to aid disease control and mitigation. |
T15 |
4778-4906 |
Epistemic_statement |
denotes |
Furthermore, knowing about the effect of environmental parameters on infection prevalence can be used for targeted intervention. |
T16 |
4907-5105 |
Epistemic_statement |
denotes |
Mathematical models can help to identify the important processes and critical points in the disease cycle and can help to evaluate the efficiency of disease management options before implementation. |
T17 |
5106-5294 |
Epistemic_statement |
denotes |
However, the testing and implementation of predictive models for aquatic diseases is challenging due to a lack of data and the complexity of host-parasite interactions (Ray et al., 2015) . |
T18 |
5886-6037 |
Epistemic_statement |
denotes |
However, neither model considers the spatial aspects of fish migration or spore concentration which should play a role in determining disease dynamics. |
T19 |
6221-6366 |
Epistemic_statement |
denotes |
As infected adult carcasses produce myxospores that re-infect the polychaete host, they represent an important link in the pathogen's life cycle. |
T20 |
6367-6492 |
Epistemic_statement |
denotes |
The potential significance of adult salmon in the infection cycle has also been mentioned by Ray (2013) and Fujiwara (2014) . |
T21 |
6493-6791 |
Epistemic_statement |
denotes |
More recently, the role of infected adult salmon in the infection cycle and the possibility of removing adult carcasses to reduce the myxospore input into the river, so as to diminishing the disease induced mortality of juvenile salmon in the following spring, has been investigated by Foott et al. |
T22 |
6801-6904 |
Epistemic_statement |
denotes |
However, they concluded that there are little effects on myxospore concentration by removing carcasses. |
T23 |
6905-7269 |
Epistemic_statement |
denotes |
In order to gain a better understanding of the myxozoan disease dynamics, including the effects of spatial migration, adult salmon, climate scenarios and dam options, we consider a complex system including dynamic processes such as hydrodynamics, environmental conditions and host-parasite interactions, where each mechanism could be studied by a model on its own. |
T24 |
7270-7483 |
Epistemic_statement |
denotes |
In an attempt to balance realism and analytical tractability, we simplify this complex system to one of three partial differential equations (PDEs) which retain, nevertheless, the dynamically important mechanisms. |
T25 |
7902-7994 |
Epistemic_statement |
denotes |
To this end, we demonstrate how our PDE model can be transformed into a dose-response model. |
T26 |
9266-9429 |
Epistemic_statement |
denotes |
Finally, we demonstrate how our model system can be reduced to a single equation, namely the dose-response model, which is applicable as a disease management tool. |
T27 |
9430-9518 |
Epistemic_statement |
denotes |
The aim of the model is to predict the prevalence of infection in the adult salmon host. |
T28 |
9705-9830 |
Epistemic_statement |
denotes |
In this section we first describe the model study area and the assumptions that are made for the spatial model (Section 2.1). |
T29 |
10122-10172 |
Epistemic_statement |
denotes |
A conceptual diagram of the model is shown in Fig. |
T30 |
11571-11678 |
Epistemic_statement |
denotes |
We assume that the zones are defined clearly, even though the transitions between the zones may be gradual. |
T31 |
12352-12448 |
Epistemic_statement |
denotes |
To reduce the complexity of the model, we approximate the river by a spatially homogeneous line. |
T32 |
12834-12941 |
Epistemic_statement |
denotes |
A positive relationship between temperature and mortality from C. shasta was first described by Udey et al. |
T33 |
13196-13351 |
Epistemic_statement |
denotes |
host distribution and density, duration/timing of host-parasite interactions), the net effect of changing environmental conditions is difficult to predict. |
T34 |
13569-13730 |
Epistemic_statement |
denotes |
Moreover, the water velocity is thought to influence the transmission rate (Bjork and Bartholomew, 2009; Ray, 2013) and the polychaete distribution additionally. |
T35 |
13731-13894 |
Epistemic_statement |
denotes |
Little is known about the influence of environmental parameters on other factors such as the polychaete population, actinospore release or the disease progression. |
T36 |
14224-14310 |
Epistemic_statement |
denotes |
The life time of actinospores in water is strongly correlated to the temperature (e.g. |
T37 |
14444-14646 |
Epistemic_statement |
denotes |
Here, for simplicity, we assume a linear relationship of life time of actinospores with temperature T. The relative decay rate δ corresponds to the inverse of the average life time and can be written as |
T38 |
14647-14787 |
Epistemic_statement |
denotes |
124 C 1 and = ν 3 have been chosen to fit the two data points mentioned above and to give integer numbers for the ease of visual appearance. |
T39 |
15301-15584 |
Epistemic_statement |
denotes |
Following Salinger and Anderson (2006) , we approximate the migration of adult salmon by a mean upstream velocity v u in − km d 1 that depends on the water temperature T and the encountered current speed v c in − km d 1 which could be expressed in terms of discharge according to Eq. |
T40 |
15944-16054 |
Epistemic_statement |
denotes |
It should be noted that the studies by Salinger and Anderson (2006) have been performed in the Columbia River. |
T41 |
16055-16232 |
Epistemic_statement |
denotes |
However, their results are similar to observations that have been made in other studies for different salmon and trout species, including both field experiments in (1) and (3)). |
T42 |
16308-16559 |
Epistemic_statement |
denotes |
Since there is no empirical formulation available describing the functional relationship of temperature, water velocity and adult salmon migration speed in the Klamath River, we here assume the same approach presented by Salinger and Anderson (2006) . |
T43 |
16880-17189 |
Epistemic_statement |
denotes |
It should be noted that we assume that not all transmitted actinospores successfully initiate an infection and we therefore distinguish between the total transmission rate β and the successful transmission rate β, which we assume to be approximately one tenth of the total transmission rate (Fujiwara, 2014) . |
T44 |
17190-17331 |
Epistemic_statement |
denotes |
To estimate the successful transmission rate β as a function of the water temperature T, we assume the following functional relationship (Eq. |
T45 |
17451-17556 |
Epistemic_statement |
denotes |
The constants and the corresponding standard deviations of the fitted parameter can be found in Table 1 . |
T46 |
17685-17859 |
Epistemic_statement |
denotes |
However, due to the relatively small effect of water velocity on the transmission rate compared to the temperature we focused only on incorporating the effect of temperature. |
T47 |
17860-18025 |
Epistemic_statement |
denotes |
Little is known about polychaete's life cycle and the influence of environmental parameters on factors such as the polychaete population or actinospore release rate. |
T48 |
18988-19079 |
Epistemic_statement |
denotes |
The spatial steady-state solution for each zone can be found analytically (see Appendix A). |
T49 |
19375-19492 |
Epistemic_statement |
denotes |
However, infected adult carcasses produce myxospores that cause infection of juvenile salmon in the following spring. |
T50 |
19493-19637 |
Epistemic_statement |
denotes |
By predicting the infection prevalence in the adult salmon host, the infection risk of juvenile salmon in the following spring can be estimated. |
T51 |
20720-20909 |
Epistemic_statement |
denotes |
Since the ratio between actinospore and myxospore particles in their model is constant, we can use outcome of this complex hydrodynamic model to validate our actinospore distribution model. |
T52 |
21416-21589 |
Epistemic_statement |
denotes |
Since the total number of infected polychaetes in each year is unknown, we ran the simulations for the three defined disease risk cases (low/medium/high polychaete numbers). |
T53 |
21637-21747 |
Epistemic_statement |
denotes |
(2016) show that an additional dam release can reduce the total spore concentration in the river by up to 50%. |
T54 |
23716-23779 |
Epistemic_statement |
denotes |
An overview of the analysed scenarios can be found in Table 2 . |
T55 |
24073-24178 |
Epistemic_statement |
denotes |
With the exponential dose-response model, the prevalence of infection POI in its general form is given by |
T56 |
24179-24286 |
Epistemic_statement |
denotes |
where k is the probability to successfully initiate a response and D the dose of the substance or pathogen. |
T57 |
24287-24459 |
Epistemic_statement |
denotes |
By using the methods of characteristics, we will show that our model system can be reduced to a single equation, which is equivalent to the exponential dose-response model. |
T58 |
24460-24592 |
Epistemic_statement |
denotes |
As mentioned above, a steady state solution for the spatial distribution of actinospores can be found analytically (see Appendix A). |
T59 |
25930-26056 |
Epistemic_statement |
denotes |
In total, the simulations of our PDE model show a very good correlation with the detailed hydrodynamic model ( = R 0.9795 2 ). |
T60 |
26787-26850 |
Epistemic_statement |
denotes |
However this trend is less pronounced than that of temperature. |
T61 |
27037-27157 |
Epistemic_statement |
denotes |
For three data points, the model shows an excellent correlation with the data points for a medium number of polychaetes. |
T62 |
27244-27297 |
Epistemic_statement |
denotes |
The results indicate that the higher the additional . |
T63 |
27387-27440 |
Epistemic_statement |
denotes |
The borders of the zones are indicated by grey lines. |
T64 |
27441-27524 |
Epistemic_statement |
denotes |
The steady state solution for each zone can be found analytically (see Appendix A). |
T65 |
28451-28568 |
Epistemic_statement |
denotes |
A dam release of = − Q s Δ 125 m 3 1 at temperatures below = ∘ T 14 C can result in a prevalence reduction up to 35%. |
T66 |
28711-28861 |
Epistemic_statement |
denotes |
Compared to the climate conditions of the years 2005 -2013, climate change is likely to increase the prevalence of infection significantly (Table 2) . |
T67 |
28951-29020 |
Epistemic_statement |
denotes |
Yet, climate change and dam removal still increase prevalence by 10%. |
T68 |
29126-29274 |
Epistemic_statement |
denotes |
Since the infection prevalence for future climate conditions is high anyhow, dam removal seems to have little quantitative effect on the prevalence. |
T69 |
29551-29624 |
Epistemic_statement |
denotes |
(6)), we can derive the exponential dose-response model (see Appendix B). |
T70 |
29625-29736 |
Epistemic_statement |
denotes |
With the mean spatial actinospore dose A over the river section we can calculate the prevalence of infection as |
T71 |
29737-29834 |
Epistemic_statement |
denotes |
With the existing empirical description of the transmission rate β Bartholomew and Foott (2010) . |
T72 |
30037-30246 |
Epistemic_statement |
denotes |
(3)) and in-situ measurement of the mean actinospore concentration A , the infection risk of the years returning salmon population can be approximated easily, without simulating the entire presented PDE model. |
T73 |
30247-30343 |
Epistemic_statement |
denotes |
Therefore, the dose-response model is suitable as a decision support tool in disease management. |
T74 |
30680-30863 |
Epistemic_statement |
denotes |
In order to aid disease control and mitigation, it is necessary to predict the impact of changing water temperatures and flow regimes, whether caused by climate change or dam removal. |
T75 |
31034-31204 |
Epistemic_statement |
denotes |
The model results and the scenario analyses provide useful information on how changing river discharge and temperature affect or can be used to affect the infection risk. |
T76 |
31527-31679 |
Epistemic_statement |
denotes |
As the actinospore distribution feeds into the infection model, this is likely to be caused by the temperature dependence of the transmission rate (Fig. |
T77 |
32092-32289 |
Epistemic_statement |
denotes |
Furthermore, we included the temperature dependence in the decay rate of the actinospores and the temperature-dependent travel time of salmon that both have not been considered by Fujiwara (2014) . |
T78 |
32290-32499 |
Epistemic_statement |
denotes |
We investigated the influence of future environmental change scenarios and showed that the prevalence of infection for future climate change scenarios in comparison to current conditions is likely to increase. |
T79 |
32597-32679 |
Epistemic_statement |
denotes |
These predictions are consistent across the climate change scenarios investigated. |
T80 |
32880-33002 |
Epistemic_statement |
denotes |
Since host-parasite dynamics are complex, the total effect of environmental change on disease dynamics is hard to predict. |
T81 |
33003-33179 |
Epistemic_statement |
denotes |
On the one hand, increasing water temperature could increase or decrease infection prevalence, depending on the current temperature and the number of infected polychaetes (Fig. |
T82 |
33185-33388 |
Epistemic_statement |
denotes |
Higher temperatures can also lead to shifts in the invertebrate host distribution (Alexander et al., 2014) or in timing and duration of spore production and host-parasite interaction (Ray et al., 2015) . |
T83 |
33389-33567 |
Epistemic_statement |
denotes |
Moreover, temperatures in Klamath River are already at the salmon's upper thermal limit (Ray et al., 2015) , which makes the future of the salmon population in the river unclear. |
T84 |
33686-33811 |
Epistemic_statement |
denotes |
In addition, varying effects related to changing temperatures will interact with changes brought about different water flows. |
T85 |
33900-34086 |
Epistemic_statement |
denotes |
On the one hand, this fragmentation is likely to increase due to a global boom in hydropower dam with at least 3700 major dams currently planned or in construction (Zarfl et al., 2015) . |
T86 |
34319-34549 |
Epistemic_statement |
denotes |
Due to its relatively small predicted effect on temperature in comparison to the impact of climate change, our model suggests that dam removal is more likely to have little influence on the predicted level of infection (Table 2) . |
T87 |
35012-35057 |
Epistemic_statement |
denotes |
However, if other factors were included (e.g. |
T88 |
35058-35264 |
Epistemic_statement |
denotes |
the influence of environmental parameters on the polychaete's life cycle or temporal and spatial variation of the host-parasite interactions) then it could be possible that the conclusions would be changed. |
T89 |
35337-35481 |
Epistemic_statement |
denotes |
In particular, dam removal not only alters water temperatures and flow, but will also change the spatial overlap between the host and parasites. |
T90 |
35482-35652 |
Epistemic_statement |
denotes |
Migration of salmon to the upper Klamath basin after dam removal could provide new hosts for existing parasite genotypes that are currently isolated upstream of the dams. |
T91 |
35653-35796 |
Epistemic_statement |
denotes |
Similarly, migrating salmon could transport new pathogenic genotypes of C. shasta to currently isolated salmon populations in the upper basin . |
T92 |
35797-35939 |
Epistemic_statement |
denotes |
In addition, the influence of environmental parameters on the polychaete's life cycle is not yet understood (Stocking and Bartholomew, 2007) . |
T93 |
35940-36087 |
Epistemic_statement |
denotes |
As we discussed before, changing environmental conditions like water temperature could lead to a range shift of the invertebrate host distribution. |
T94 |
36088-36195 |
Epistemic_statement |
denotes |
The polychaete density P 0 or the release rate κ, for example, could be functions of the water temperature. |
T95 |
36196-36348 |
Epistemic_statement |
denotes |
It has been shown elsewhere that flow manipulation can be used for reducing the polychaete density (Alexander et al., 2014) or the spore concentration . |
T96 |
36349-36529 |
Epistemic_statement |
denotes |
Other management options, as for instance removing adult carcasses to reduce the myxospore input (Foott et al., 2016) , have been shown to have little effect on the infection risk. |
T97 |
36530-36751 |
Epistemic_statement |
denotes |
Our model results show that dam release before the salmon run can reduce the actinospore concentration as well as the infection prevalence in returning salmon and can therefore be a useful tool for managing Ceratomyxosis. |
T98 |
36752-36944 |
Epistemic_statement |
denotes |
We showed that our model, although based on a number of mechanistic subprocesses, can be simplified to an exponential dose-response function that can be integrated in decision support systems. |
T99 |
36945-37083 |
Epistemic_statement |
denotes |
Despite the simplicity of this function, it encapsulates all significant mechanisms of the disease dynamics in adult fish considered here. |
T100 |
37620-37824 |
Epistemic_statement |
denotes |
Discharge, temperature and spore concentration are easy to measure and can be used to estimate the infection risk with the dose-response model and evaluate the effectiveness of disease management options. |
T101 |
38979-39056 |
Epistemic_statement |
denotes |
We assume the temperature T and the water velocity v c to be constant so that |
T102 |
39057-39085 |
Epistemic_statement |
denotes |
In a first step we solve Eq. |