| | | | | | | | | | | | |
|---|
| A | 185 | Produce CBFish Status Report | Periodic Status Reports for BPA | The Contractor shall report on the status of milestones and deliverables in Pisces. Reports shall be completed either monthly or quarterly as determined by the BPA COTR. Additionally, when indicating a deliverable milestone as COMPLETE, the contractor shall provide metrics and the final location (latitude and longitude) prior to submitting the report to the BPA COTR. | A | SR | Oct-Dec 2010 (10/1/2010 - 12/31/2010) | 01/01/2011 | 01/15/2011 | Concluded | Richard Golden Jr (Inactive) | 07/20/2010 1:32 PM |
| A | 185 | Produce CBFish Status Report | Periodic Status Reports for BPA | The Contractor shall report on the status of milestones and deliverables in Pisces. Reports shall be completed either monthly or quarterly as determined by the BPA COTR. Additionally, when indicating a deliverable milestone as COMPLETE, the contractor shall provide metrics and the final location (latitude and longitude) prior to submitting the report to the BPA COTR. | B | SR | Jan-Mar 2011 (1/1/2011 - 3/31/2011) | 04/01/2011 | 04/15/2011 | Concluded | Richard Golden Jr (Inactive) | 07/20/2010 1:32 PM |
| A | 185 | Produce CBFish Status Report | Periodic Status Reports for BPA | The Contractor shall report on the status of milestones and deliverables in Pisces. Reports shall be completed either monthly or quarterly as determined by the BPA COTR. Additionally, when indicating a deliverable milestone as COMPLETE, the contractor shall provide metrics and the final location (latitude and longitude) prior to submitting the report to the BPA COTR. | C | SR | Apr-Jun 2011 (4/1/2011 - 6/30/2011) | 07/01/2011 | 07/15/2011 | Concluded | Richard Golden Jr (Inactive) | 07/20/2010 1:32 PM |
| A | 185 | Produce CBFish Status Report | Periodic Status Reports for BPA | The Contractor shall report on the status of milestones and deliverables in Pisces. Reports shall be completed either monthly or quarterly as determined by the BPA COTR. Additionally, when indicating a deliverable milestone as COMPLETE, the contractor shall provide metrics and the final location (latitude and longitude) prior to submitting the report to the BPA COTR. | D | SR | Final Jul-Sep 2011 (7/1/2011 - 9/30/2011) | 09/16/2011 | 09/30/2011 | Concluded | Richard Golden Jr (Inactive) | 07/20/2010 1:32 PM |
| B | 165 | Produce Environmental Compliance Documentation | Categorical Exclusion | Categorical Exclusion Applied (from Subpart D, 10 C.F.R. Part 1021): B3.3: Field and laboratory research, inventory, and information collection activities that are directly related to the conservation of fish or wildlife resources and that involve only negligible habitat destruction or population reduction. | A | DELIV | Complies with NEPA | | 10/01/2010 | Concluded | Michael Blouin | 07/19/2010 12:12 PM |
| C | 157 | Collect/Generate/Validate Field and Lab Data | Genotyping | Summarize statistics for fish genotyped to date, give progress assessment. Approximately 750 new fish genotyped by end of each reporting quarter. More details on loci and methods are explained in Araki et al., (2006; Conservation Biology, 21:181-190). | A | EC | Environmental compliance requirements complete | 10/01/2010 | 07/19/2011 | Concluded | Michael Blouin | 07/20/2010 11:05 AM |
| C | 157 | Collect/Generate/Validate Field and Lab Data | Genotyping | Summarize statistics for fish genotyped to date, give progress assessment. Approximately 750 new fish genotyped by end of each reporting quarter. More details on loci and methods are explained in Araki et al., (2006; Conservation Biology, 21:181-190). | B | | Approximately 750 fish genotyped | 10/01/2010 | 12/28/2010 | Concluded | Michael Blouin | 07/19/2010 12:19 PM |
| C | 157 | Collect/Generate/Validate Field and Lab Data | Genotyping | Summarize statistics for fish genotyped to date, give progress assessment. Approximately 750 new fish genotyped by end of each reporting quarter. More details on loci and methods are explained in Araki et al., (2006; Conservation Biology, 21:181-190). | C | | Approximately 750 fish genotyped | 01/01/2011 | 03/31/2011 | Concluded | Michael Blouin | 07/19/2010 12:19 PM |
| C | 157 | Collect/Generate/Validate Field and Lab Data | Genotyping | Summarize statistics for fish genotyped to date, give progress assessment. Approximately 750 new fish genotyped by end of each reporting quarter. More details on loci and methods are explained in Araki et al., (2006; Conservation Biology, 21:181-190). | D | | Approximately 750 fish genotyped | 04/01/2011 | 06/30/2011 | Concluded | Michael Blouin | 07/19/2010 12:19 PM |
| C | 157 | Collect/Generate/Validate Field and Lab Data | Genotyping | Summarize statistics for fish genotyped to date, give progress assessment. Approximately 750 new fish genotyped by end of each reporting quarter. More details on loci and methods are explained in Araki et al., (2006; Conservation Biology, 21:181-190). | E | | Approximately 750 fish genotyped | 07/01/2011 | 09/30/2011 | Concluded | Michael Blouin | 07/19/2010 12:19 PM |
| C | 157 | Collect/Generate/Validate Field and Lab Data | Genotyping | Summarize statistics for fish genotyped to date, give progress assessment. Approximately 750 new fish genotyped by end of each reporting quarter. More details on loci and methods are explained in Araki et al., (2006; Conservation Biology, 21:181-190). | F | DELIV | Successfully genotype approximately 3000 fish | | 09/30/2011 | Concluded | Michael Blouin | 07/19/2010 12:14 PM |
| D | 162 | Analyze/Interpret Data | Data Analysis | Our data analysis efforts over the next year will focus on the following three actions:
(1) Estimate fitness of summer-run Hnew fish relative to that of wild summer-run
It is important to know whether our results with the winter-run steelhead are a special case, or are typical of hatchery supplementation programs. Several other research groups are in the process of examining the relative fitness of first-generation hatchery salmon of various species (coho, Chinook, steelhead), so eventually we will have a basis for comparison. But these studies have not published their results yet. We can now analyze the relative fitness of the first-generation summer-run steelhead stock that were released into the Hood River beginning in 1997. Those hatchery fish began returning in appreciable numbers in 2000. Because the winter and summer run are reproductively independent and breed in different parts of the Hood River, these summer-run data should constitute an independent test of the hypothesis that first-generation hatchery steelhead have lower fitness than wild fish. As part of the FY 2010 objectives we have been analyzing the first two run years (2000 and 2001) in which these summer-run Hnew bred in the wild (for these two years we should now have in hand >95% of their returning offspring). We will continue with the next year, with the ultimate goal of having four years of comparisons between Hnew summers and wild fish.
(2) Test whether offspring of hatchery fish grow faster under hatchery conditions than the offspring of wild fish
The 30-40% per generation drop in fitness that we estimated is occurring per hatchery generation in the winter run (Araki et al., 2007d) is so extreme that it was questionable whether domestication selection was even a plausible explanation. Here “domestication selection” means both positive selection for traits that are favored in the hatchery and relaxation of natural selection. We did a quantitative genetic analysis of selection in alternating environments and showed that strong selection was indeed a sufficient explanation for our results (Araki et al., 2008). So now the question becomes, what traits are involved?
Both adult traits and juvenile traits could be the targets of positive selection or relaxed natural selection. Artificial spawning in the hatchery almost certainly results in relaxation of mate selection (Berejikian et al.2000; de Gaudemar et al. 2000), intra-sexual competition, and selection on traits such as body size, egg size, fecundity and spawn timing and location (van den Bergheand Gross 1989; Fleming and Gross 1994; Einum and Fleming 2000). As for juvenile traits, given only 5-15% mortality in hatcheries, there is not enough selective death to generate the fitness declines observed. So the effect cannot be explained simply by viability (survival) selection on some trait in the hatchery. On the other hand, mortality from smolt to adult is often > 99%. So juvenile traits that affect adult survival (or perhaps even breeding behaviors) could well be the targets of selection. For example, smolt size is positively correlated with ocean survival, so there is strong selection on size at release (Reisenbichler et al., 2004). In the high-food and predator-free hatchery environment, this survival difference should select for high growth rate in hatchery juveniles, perhaps via behavioral or metabolic changes. But an excessively high growth rate can be maladaptive in natural environments (Arendt, 1997). So when those surviving hatchery fish reproduce in a natural setting, their offspring could have lower survival than wild offspring owing to inappropriate behaviors, an excessively high metabolism, or other maladaptive consequences of a genetic predisposition for fast growth in the benign and high-food hatchery environment (Sundstrom et al., 2005; Biro et al., 2004; Tymchuk et al., 2007). Therefore, we have begun testing whether there is evidence that the Hood River hatchery fish have been selected to have higher growth rates under hatchery conditions. In 2008 we set up a series of 2x2 matrix crosses involving a hatchery and wild male by a hatchery and wild female, and then raised each pooled group of four families in the hatchery for seven weeks. We measured the size of each fish and archived a tissue sample for DNA typing. After sorting them back into their respective families, we found no evidence for an additive genetic effect of hatchery versus wild background, although there was a weak maternal effect in which the offspring of wild females seemed to grow faster (perhaps an effect of egg size?). We started a second set of crosses in 2009 and sampled them at 2 months of age and one year of age. Among the 2-month olds we again found little evidence for a difference between fish of H vs. W backgrounds. We now have their siblings raised to one year of age, and plan to analyze them the same way. It is possible that any growth difference will appear later in life, so we haven’t rule out the growth rate hypothesis yet. Finally, we are setting up a 3rd set of crosses this summer, and will replicate the experiment in one final year, although raising them at densities that more closely match those of the production fish (last year’s fish were not as crowded). If we still see no effect of genetic background, then we will consider the growth rate hypothesis to be not so compelling.
(3) Test of differential disease resistance:
In collaboration with Jeri Bartholomew at OSU, we will attempt to test the siblings of this year’s crosses with viral and bacterial challenges. These experiments are to be conducted at the salmon disease lab at OSU. The fish will be challenged in groups and we will sort them back into their respective families after the fact via microsatellites, as with the growth rate study. Our hypothesis is that fish of H background will be less disease resistant as a consequence of hatchery selection for growth (i.e. selection to invest energetic resources into growth at the expense of maintenance). We haven't done this before, so we aren't guaranteeing that this first year's experiment will work (e.g. we have to estimate the proper challenge doses, fish can die for other reasons, and so on). But if it does work, and we do find a difference, it will be a very interesting corroborating evidence for selection on key life history traits as the mechanism of domestication.
References cited:
Araki, H., W.R. Ardren, E. Olsen, B. Cooper and M.S. Blouin. 2007a. Reproductive success of captive-bred steelhead trout in the wild: evaluation of three hatchery programs in the Hood River. Conservation Biology 21:181-190.
Araki, H., B. Cooper and M.S. Blouin. 2007d. Genetic effects of captive breeding cause a rapid, cumulative fitness decline in the wild. Science 318: 100-103.
Araki, H., B. Berejikian, M. Ford, and M.S. Blouin. 2008 Fitness of hatchery-reared salmonids in the wild. Evolutionary Applications 1:342-355.
Arendt, J.D.. 1997. Adaptive intrinsic growth rates: an integration across taxa. Quarterly Review of Biology 72:149-177.
Berejikian, B. A., E. P. Tezak, and A. L. LaRae. 2000. Female mate choice and spawning behavior of chinook salmon under experimental conditions. Journal of Fish Biology 57:647–661.
van den Berghe, E. P., and M. R. Gross. 1989. Natural selection resulting from female breeding competition in a Pacific salmon (Coho: Oncorhynchus kisutch). Evolution 43:125–140
Biro, P. A., M. V. Abrahams, J. R. Post, and E. A. Parkinson. 2004. Predators select against high growth rates and risk-taking behaviour in domestic trout populations. Proceedings: Biological Sciences 271:2233.
Einum, S., and I. A. Fleming. 2000. Selection against late emergence and small offspring in Atlantic salmon (Salmo salar). Evolution 54:628–639.
Fleming, I. A., and M. R. Gross. 1994. Breeding success of hatchery and wild coho salmon (Oncorhynchus kisutch) in competition. Ecological Applications 3:230–245.
de Gaudemar, B., J. M. Bonzom, and E. Beall. 2000. Effects of courtship and relative mate size on sexual motivation in Atlantic salmon. Journal of Fish Biology 57:502–515.
Reisenbichler, R., S. Rubin, L. Wetzel, and S. Phelps. 2004. Natural selection after release from a hatchery leads to domestication in steelhead, Oncorhynchus mykiss. Pp. 371-383 in B. R. Howell, E. Moksness and T. Svåsand, eds. Stock enhancement and sea ranching. Oxford, Malden, MA.
Seamons, T. R., P. Bentzen, and T. P. Quinn. 2004. The mating system of steelhead, Oncorhynchus mykiss, inferred by molecular analysis of parents and progeny. Environmental Biology of Fishes 69: 333–344
Sundstrom, L. F., M. Lohmus, R. H. Devlin, and E. Brainerd. 2005. Selection On Increased Intrinsic Growth Rates In Coho Salmon, Oncorhynchus Kisutch. Evolution 59:1560.
Tymchuk, W. E., L. F. Sundstrom, and R. H. Devlin. 2007. Growth And Survival Trade-Offs And Outbreeding Depression In Rainbow Trout (Oncorhynchus Mykiss). Evolution 61:1225.
.Zimmerman, C. E., and G. H. Reeves. 2000. Population structure of sympatric anadromous and nonanadromous Oncorhynchusmykiss: evidence from spawning surveys and otolith microchemistry. Canadian Journal of Fisheries and Aquatic Sciences 57:2152–2162 | A | | Begin interpretation of genotype data | 07/01/2011 | 09/30/2011 | Concluded | Michael Blouin | 07/19/2010 12:21 PM |
| D | 162 | Analyze/Interpret Data | Data Analysis | Our data analysis efforts over the next year will focus on the following three actions:
(1) Estimate fitness of summer-run Hnew fish relative to that of wild summer-run
It is important to know whether our results with the winter-run steelhead are a special case, or are typical of hatchery supplementation programs. Several other research groups are in the process of examining the relative fitness of first-generation hatchery salmon of various species (coho, Chinook, steelhead), so eventually we will have a basis for comparison. But these studies have not published their results yet. We can now analyze the relative fitness of the first-generation summer-run steelhead stock that were released into the Hood River beginning in 1997. Those hatchery fish began returning in appreciable numbers in 2000. Because the winter and summer run are reproductively independent and breed in different parts of the Hood River, these summer-run data should constitute an independent test of the hypothesis that first-generation hatchery steelhead have lower fitness than wild fish. As part of the FY 2010 objectives we have been analyzing the first two run years (2000 and 2001) in which these summer-run Hnew bred in the wild (for these two years we should now have in hand >95% of their returning offspring). We will continue with the next year, with the ultimate goal of having four years of comparisons between Hnew summers and wild fish.
(2) Test whether offspring of hatchery fish grow faster under hatchery conditions than the offspring of wild fish
The 30-40% per generation drop in fitness that we estimated is occurring per hatchery generation in the winter run (Araki et al., 2007d) is so extreme that it was questionable whether domestication selection was even a plausible explanation. Here “domestication selection” means both positive selection for traits that are favored in the hatchery and relaxation of natural selection. We did a quantitative genetic analysis of selection in alternating environments and showed that strong selection was indeed a sufficient explanation for our results (Araki et al., 2008). So now the question becomes, what traits are involved?
Both adult traits and juvenile traits could be the targets of positive selection or relaxed natural selection. Artificial spawning in the hatchery almost certainly results in relaxation of mate selection (Berejikian et al.2000; de Gaudemar et al. 2000), intra-sexual competition, and selection on traits such as body size, egg size, fecundity and spawn timing and location (van den Bergheand Gross 1989; Fleming and Gross 1994; Einum and Fleming 2000). As for juvenile traits, given only 5-15% mortality in hatcheries, there is not enough selective death to generate the fitness declines observed. So the effect cannot be explained simply by viability (survival) selection on some trait in the hatchery. On the other hand, mortality from smolt to adult is often > 99%. So juvenile traits that affect adult survival (or perhaps even breeding behaviors) could well be the targets of selection. For example, smolt size is positively correlated with ocean survival, so there is strong selection on size at release (Reisenbichler et al., 2004). In the high-food and predator-free hatchery environment, this survival difference should select for high growth rate in hatchery juveniles, perhaps via behavioral or metabolic changes. But an excessively high growth rate can be maladaptive in natural environments (Arendt, 1997). So when those surviving hatchery fish reproduce in a natural setting, their offspring could have lower survival than wild offspring owing to inappropriate behaviors, an excessively high metabolism, or other maladaptive consequences of a genetic predisposition for fast growth in the benign and high-food hatchery environment (Sundstrom et al., 2005; Biro et al., 2004; Tymchuk et al., 2007). Therefore, we have begun testing whether there is evidence that the Hood River hatchery fish have been selected to have higher growth rates under hatchery conditions. In 2008 we set up a series of 2x2 matrix crosses involving a hatchery and wild male by a hatchery and wild female, and then raised each pooled group of four families in the hatchery for seven weeks. We measured the size of each fish and archived a tissue sample for DNA typing. After sorting them back into their respective families, we found no evidence for an additive genetic effect of hatchery versus wild background, although there was a weak maternal effect in which the offspring of wild females seemed to grow faster (perhaps an effect of egg size?). We started a second set of crosses in 2009 and sampled them at 2 months of age and one year of age. Among the 2-month olds we again found little evidence for a difference between fish of H vs. W backgrounds. We now have their siblings raised to one year of age, and plan to analyze them the same way. It is possible that any growth difference will appear later in life, so we haven’t rule out the growth rate hypothesis yet. Finally, we are setting up a 3rd set of crosses this summer, and will replicate the experiment in one final year, although raising them at densities that more closely match those of the production fish (last year’s fish were not as crowded). If we still see no effect of genetic background, then we will consider the growth rate hypothesis to be not so compelling.
(3) Test of differential disease resistance:
In collaboration with Jeri Bartholomew at OSU, we will attempt to test the siblings of this year’s crosses with viral and bacterial challenges. These experiments are to be conducted at the salmon disease lab at OSU. The fish will be challenged in groups and we will sort them back into their respective families after the fact via microsatellites, as with the growth rate study. Our hypothesis is that fish of H background will be less disease resistant as a consequence of hatchery selection for growth (i.e. selection to invest energetic resources into growth at the expense of maintenance). We haven't done this before, so we aren't guaranteeing that this first year's experiment will work (e.g. we have to estimate the proper challenge doses, fish can die for other reasons, and so on). But if it does work, and we do find a difference, it will be a very interesting corroborating evidence for selection on key life history traits as the mechanism of domestication.
References cited:
Araki, H., W.R. Ardren, E. Olsen, B. Cooper and M.S. Blouin. 2007a. Reproductive success of captive-bred steelhead trout in the wild: evaluation of three hatchery programs in the Hood River. Conservation Biology 21:181-190.
Araki, H., B. Cooper and M.S. Blouin. 2007d. Genetic effects of captive breeding cause a rapid, cumulative fitness decline in the wild. Science 318: 100-103.
Araki, H., B. Berejikian, M. Ford, and M.S. Blouin. 2008 Fitness of hatchery-reared salmonids in the wild. Evolutionary Applications 1:342-355.
Arendt, J.D.. 1997. Adaptive intrinsic growth rates: an integration across taxa. Quarterly Review of Biology 72:149-177.
Berejikian, B. A., E. P. Tezak, and A. L. LaRae. 2000. Female mate choice and spawning behavior of chinook salmon under experimental conditions. Journal of Fish Biology 57:647–661.
van den Berghe, E. P., and M. R. Gross. 1989. Natural selection resulting from female breeding competition in a Pacific salmon (Coho: Oncorhynchus kisutch). Evolution 43:125–140
Biro, P. A., M. V. Abrahams, J. R. Post, and E. A. Parkinson. 2004. Predators select against high growth rates and risk-taking behaviour in domestic trout populations. Proceedings: Biological Sciences 271:2233.
Einum, S., and I. A. Fleming. 2000. Selection against late emergence and small offspring in Atlantic salmon (Salmo salar). Evolution 54:628–639.
Fleming, I. A., and M. R. Gross. 1994. Breeding success of hatchery and wild coho salmon (Oncorhynchus kisutch) in competition. Ecological Applications 3:230–245.
de Gaudemar, B., J. M. Bonzom, and E. Beall. 2000. Effects of courtship and relative mate size on sexual motivation in Atlantic salmon. Journal of Fish Biology 57:502–515.
Reisenbichler, R., S. Rubin, L. Wetzel, and S. Phelps. 2004. Natural selection after release from a hatchery leads to domestication in steelhead, Oncorhynchus mykiss. Pp. 371-383 in B. R. Howell, E. Moksness and T. Svåsand, eds. Stock enhancement and sea ranching. Oxford, Malden, MA.
Seamons, T. R., P. Bentzen, and T. P. Quinn. 2004. The mating system of steelhead, Oncorhynchus mykiss, inferred by molecular analysis of parents and progeny. Environmental Biology of Fishes 69: 333–344
Sundstrom, L. F., M. Lohmus, R. H. Devlin, and E. Brainerd. 2005. Selection On Increased Intrinsic Growth Rates In Coho Salmon, Oncorhynchus Kisutch. Evolution 59:1560.
Tymchuk, W. E., L. F. Sundstrom, and R. H. Devlin. 2007. Growth And Survival Trade-Offs And Outbreeding Depression In Rainbow Trout (Oncorhynchus Mykiss). Evolution 61:1225.
.Zimmerman, C. E., and G. H. Reeves. 2000. Population structure of sympatric anadromous and nonanadromous Oncorhynchusmykiss: evidence from spawning surveys and otolith microchemistry. Canadian Journal of Fisheries and Aquatic Sciences 57:2152–2162 | B | | Estimate fitness of summer-run fish | 07/01/2011 | 09/30/2011 | Concluded | Michael Blouin | 07/19/2010 12:21 PM |
| D | 162 | Analyze/Interpret Data | Data Analysis | Our data analysis efforts over the next year will focus on the following three actions:
(1) Estimate fitness of summer-run Hnew fish relative to that of wild summer-run
It is important to know whether our results with the winter-run steelhead are a special case, or are typical of hatchery supplementation programs. Several other research groups are in the process of examining the relative fitness of first-generation hatchery salmon of various species (coho, Chinook, steelhead), so eventually we will have a basis for comparison. But these studies have not published their results yet. We can now analyze the relative fitness of the first-generation summer-run steelhead stock that were released into the Hood River beginning in 1997. Those hatchery fish began returning in appreciable numbers in 2000. Because the winter and summer run are reproductively independent and breed in different parts of the Hood River, these summer-run data should constitute an independent test of the hypothesis that first-generation hatchery steelhead have lower fitness than wild fish. As part of the FY 2010 objectives we have been analyzing the first two run years (2000 and 2001) in which these summer-run Hnew bred in the wild (for these two years we should now have in hand >95% of their returning offspring). We will continue with the next year, with the ultimate goal of having four years of comparisons between Hnew summers and wild fish.
(2) Test whether offspring of hatchery fish grow faster under hatchery conditions than the offspring of wild fish
The 30-40% per generation drop in fitness that we estimated is occurring per hatchery generation in the winter run (Araki et al., 2007d) is so extreme that it was questionable whether domestication selection was even a plausible explanation. Here “domestication selection” means both positive selection for traits that are favored in the hatchery and relaxation of natural selection. We did a quantitative genetic analysis of selection in alternating environments and showed that strong selection was indeed a sufficient explanation for our results (Araki et al., 2008). So now the question becomes, what traits are involved?
Both adult traits and juvenile traits could be the targets of positive selection or relaxed natural selection. Artificial spawning in the hatchery almost certainly results in relaxation of mate selection (Berejikian et al.2000; de Gaudemar et al. 2000), intra-sexual competition, and selection on traits such as body size, egg size, fecundity and spawn timing and location (van den Bergheand Gross 1989; Fleming and Gross 1994; Einum and Fleming 2000). As for juvenile traits, given only 5-15% mortality in hatcheries, there is not enough selective death to generate the fitness declines observed. So the effect cannot be explained simply by viability (survival) selection on some trait in the hatchery. On the other hand, mortality from smolt to adult is often > 99%. So juvenile traits that affect adult survival (or perhaps even breeding behaviors) could well be the targets of selection. For example, smolt size is positively correlated with ocean survival, so there is strong selection on size at release (Reisenbichler et al., 2004). In the high-food and predator-free hatchery environment, this survival difference should select for high growth rate in hatchery juveniles, perhaps via behavioral or metabolic changes. But an excessively high growth rate can be maladaptive in natural environments (Arendt, 1997). So when those surviving hatchery fish reproduce in a natural setting, their offspring could have lower survival than wild offspring owing to inappropriate behaviors, an excessively high metabolism, or other maladaptive consequences of a genetic predisposition for fast growth in the benign and high-food hatchery environment (Sundstrom et al., 2005; Biro et al., 2004; Tymchuk et al., 2007). Therefore, we have begun testing whether there is evidence that the Hood River hatchery fish have been selected to have higher growth rates under hatchery conditions. In 2008 we set up a series of 2x2 matrix crosses involving a hatchery and wild male by a hatchery and wild female, and then raised each pooled group of four families in the hatchery for seven weeks. We measured the size of each fish and archived a tissue sample for DNA typing. After sorting them back into their respective families, we found no evidence for an additive genetic effect of hatchery versus wild background, although there was a weak maternal effect in which the offspring of wild females seemed to grow faster (perhaps an effect of egg size?). We started a second set of crosses in 2009 and sampled them at 2 months of age and one year of age. Among the 2-month olds we again found little evidence for a difference between fish of H vs. W backgrounds. We now have their siblings raised to one year of age, and plan to analyze them the same way. It is possible that any growth difference will appear later in life, so we haven’t rule out the growth rate hypothesis yet. Finally, we are setting up a 3rd set of crosses this summer, and will replicate the experiment in one final year, although raising them at densities that more closely match those of the production fish (last year’s fish were not as crowded). If we still see no effect of genetic background, then we will consider the growth rate hypothesis to be not so compelling.
(3) Test of differential disease resistance:
In collaboration with Jeri Bartholomew at OSU, we will attempt to test the siblings of this year’s crosses with viral and bacterial challenges. These experiments are to be conducted at the salmon disease lab at OSU. The fish will be challenged in groups and we will sort them back into their respective families after the fact via microsatellites, as with the growth rate study. Our hypothesis is that fish of H background will be less disease resistant as a consequence of hatchery selection for growth (i.e. selection to invest energetic resources into growth at the expense of maintenance). We haven't done this before, so we aren't guaranteeing that this first year's experiment will work (e.g. we have to estimate the proper challenge doses, fish can die for other reasons, and so on). But if it does work, and we do find a difference, it will be a very interesting corroborating evidence for selection on key life history traits as the mechanism of domestication.
References cited:
Araki, H., W.R. Ardren, E. Olsen, B. Cooper and M.S. Blouin. 2007a. Reproductive success of captive-bred steelhead trout in the wild: evaluation of three hatchery programs in the Hood River. Conservation Biology 21:181-190.
Araki, H., B. Cooper and M.S. Blouin. 2007d. Genetic effects of captive breeding cause a rapid, cumulative fitness decline in the wild. Science 318: 100-103.
Araki, H., B. Berejikian, M. Ford, and M.S. Blouin. 2008 Fitness of hatchery-reared salmonids in the wild. Evolutionary Applications 1:342-355.
Arendt, J.D.. 1997. Adaptive intrinsic growth rates: an integration across taxa. Quarterly Review of Biology 72:149-177.
Berejikian, B. A., E. P. Tezak, and A. L. LaRae. 2000. Female mate choice and spawning behavior of chinook salmon under experimental conditions. Journal of Fish Biology 57:647–661.
van den Berghe, E. P., and M. R. Gross. 1989. Natural selection resulting from female breeding competition in a Pacific salmon (Coho: Oncorhynchus kisutch). Evolution 43:125–140
Biro, P. A., M. V. Abrahams, J. R. Post, and E. A. Parkinson. 2004. Predators select against high growth rates and risk-taking behaviour in domestic trout populations. Proceedings: Biological Sciences 271:2233.
Einum, S., and I. A. Fleming. 2000. Selection against late emergence and small offspring in Atlantic salmon (Salmo salar). Evolution 54:628–639.
Fleming, I. A., and M. R. Gross. 1994. Breeding success of hatchery and wild coho salmon (Oncorhynchus kisutch) in competition. Ecological Applications 3:230–245.
de Gaudemar, B., J. M. Bonzom, and E. Beall. 2000. Effects of courtship and relative mate size on sexual motivation in Atlantic salmon. Journal of Fish Biology 57:502–515.
Reisenbichler, R., S. Rubin, L. Wetzel, and S. Phelps. 2004. Natural selection after release from a hatchery leads to domestication in steelhead, Oncorhynchus mykiss. Pp. 371-383 in B. R. Howell, E. Moksness and T. Svåsand, eds. Stock enhancement and sea ranching. Oxford, Malden, MA.
Seamons, T. R., P. Bentzen, and T. P. Quinn. 2004. The mating system of steelhead, Oncorhynchus mykiss, inferred by molecular analysis of parents and progeny. Environmental Biology of Fishes 69: 333–344
Sundstrom, L. F., M. Lohmus, R. H. Devlin, and E. Brainerd. 2005. Selection On Increased Intrinsic Growth Rates In Coho Salmon, Oncorhynchus Kisutch. Evolution 59:1560.
Tymchuk, W. E., L. F. Sundstrom, and R. H. Devlin. 2007. Growth And Survival Trade-Offs And Outbreeding Depression In Rainbow Trout (Oncorhynchus Mykiss). Evolution 61:1225.
.Zimmerman, C. E., and G. H. Reeves. 2000. Population structure of sympatric anadromous and nonanadromous Oncorhynchusmykiss: evidence from spawning surveys and otolith microchemistry. Canadian Journal of Fisheries and Aquatic Sciences 57:2152–2162 | C | | Compare growth | 07/01/2011 | 09/30/2011 | Concluded | Michael Blouin | 07/19/2010 12:21 PM |
| D | 162 | Analyze/Interpret Data | Data Analysis | Our data analysis efforts over the next year will focus on the following three actions:
(1) Estimate fitness of summer-run Hnew fish relative to that of wild summer-run
It is important to know whether our results with the winter-run steelhead are a special case, or are typical of hatchery supplementation programs. Several other research groups are in the process of examining the relative fitness of first-generation hatchery salmon of various species (coho, Chinook, steelhead), so eventually we will have a basis for comparison. But these studies have not published their results yet. We can now analyze the relative fitness of the first-generation summer-run steelhead stock that were released into the Hood River beginning in 1997. Those hatchery fish began returning in appreciable numbers in 2000. Because the winter and summer run are reproductively independent and breed in different parts of the Hood River, these summer-run data should constitute an independent test of the hypothesis that first-generation hatchery steelhead have lower fitness than wild fish. As part of the FY 2010 objectives we have been analyzing the first two run years (2000 and 2001) in which these summer-run Hnew bred in the wild (for these two years we should now have in hand >95% of their returning offspring). We will continue with the next year, with the ultimate goal of having four years of comparisons between Hnew summers and wild fish.
(2) Test whether offspring of hatchery fish grow faster under hatchery conditions than the offspring of wild fish
The 30-40% per generation drop in fitness that we estimated is occurring per hatchery generation in the winter run (Araki et al., 2007d) is so extreme that it was questionable whether domestication selection was even a plausible explanation. Here “domestication selection” means both positive selection for traits that are favored in the hatchery and relaxation of natural selection. We did a quantitative genetic analysis of selection in alternating environments and showed that strong selection was indeed a sufficient explanation for our results (Araki et al., 2008). So now the question becomes, what traits are involved?
Both adult traits and juvenile traits could be the targets of positive selection or relaxed natural selection. Artificial spawning in the hatchery almost certainly results in relaxation of mate selection (Berejikian et al.2000; de Gaudemar et al. 2000), intra-sexual competition, and selection on traits such as body size, egg size, fecundity and spawn timing and location (van den Bergheand Gross 1989; Fleming and Gross 1994; Einum and Fleming 2000). As for juvenile traits, given only 5-15% mortality in hatcheries, there is not enough selective death to generate the fitness declines observed. So the effect cannot be explained simply by viability (survival) selection on some trait in the hatchery. On the other hand, mortality from smolt to adult is often > 99%. So juvenile traits that affect adult survival (or perhaps even breeding behaviors) could well be the targets of selection. For example, smolt size is positively correlated with ocean survival, so there is strong selection on size at release (Reisenbichler et al., 2004). In the high-food and predator-free hatchery environment, this survival difference should select for high growth rate in hatchery juveniles, perhaps via behavioral or metabolic changes. But an excessively high growth rate can be maladaptive in natural environments (Arendt, 1997). So when those surviving hatchery fish reproduce in a natural setting, their offspring could have lower survival than wild offspring owing to inappropriate behaviors, an excessively high metabolism, or other maladaptive consequences of a genetic predisposition for fast growth in the benign and high-food hatchery environment (Sundstrom et al., 2005; Biro et al., 2004; Tymchuk et al., 2007). Therefore, we have begun testing whether there is evidence that the Hood River hatchery fish have been selected to have higher growth rates under hatchery conditions. In 2008 we set up a series of 2x2 matrix crosses involving a hatchery and wild male by a hatchery and wild female, and then raised each pooled group of four families in the hatchery for seven weeks. We measured the size of each fish and archived a tissue sample for DNA typing. After sorting them back into their respective families, we found no evidence for an additive genetic effect of hatchery versus wild background, although there was a weak maternal effect in which the offspring of wild females seemed to grow faster (perhaps an effect of egg size?). We started a second set of crosses in 2009 and sampled them at 2 months of age and one year of age. Among the 2-month olds we again found little evidence for a difference between fish of H vs. W backgrounds. We now have their siblings raised to one year of age, and plan to analyze them the same way. It is possible that any growth difference will appear later in life, so we haven’t rule out the growth rate hypothesis yet. Finally, we are setting up a 3rd set of crosses this summer, and will replicate the experiment in one final year, although raising them at densities that more closely match those of the production fish (last year’s fish were not as crowded). If we still see no effect of genetic background, then we will consider the growth rate hypothesis to be not so compelling.
(3) Test of differential disease resistance:
In collaboration with Jeri Bartholomew at OSU, we will attempt to test the siblings of this year’s crosses with viral and bacterial challenges. These experiments are to be conducted at the salmon disease lab at OSU. The fish will be challenged in groups and we will sort them back into their respective families after the fact via microsatellites, as with the growth rate study. Our hypothesis is that fish of H background will be less disease resistant as a consequence of hatchery selection for growth (i.e. selection to invest energetic resources into growth at the expense of maintenance). We haven't done this before, so we aren't guaranteeing that this first year's experiment will work (e.g. we have to estimate the proper challenge doses, fish can die for other reasons, and so on). But if it does work, and we do find a difference, it will be a very interesting corroborating evidence for selection on key life history traits as the mechanism of domestication.
References cited:
Araki, H., W.R. Ardren, E. Olsen, B. Cooper and M.S. Blouin. 2007a. Reproductive success of captive-bred steelhead trout in the wild: evaluation of three hatchery programs in the Hood River. Conservation Biology 21:181-190.
Araki, H., B. Cooper and M.S. Blouin. 2007d. Genetic effects of captive breeding cause a rapid, cumulative fitness decline in the wild. Science 318: 100-103.
Araki, H., B. Berejikian, M. Ford, and M.S. Blouin. 2008 Fitness of hatchery-reared salmonids in the wild. Evolutionary Applications 1:342-355.
Arendt, J.D.. 1997. Adaptive intrinsic growth rates: an integration across taxa. Quarterly Review of Biology 72:149-177.
Berejikian, B. A., E. P. Tezak, and A. L. LaRae. 2000. Female mate choice and spawning behavior of chinook salmon under experimental conditions. Journal of Fish Biology 57:647–661.
van den Berghe, E. P., and M. R. Gross. 1989. Natural selection resulting from female breeding competition in a Pacific salmon (Coho: Oncorhynchus kisutch). Evolution 43:125–140
Biro, P. A., M. V. Abrahams, J. R. Post, and E. A. Parkinson. 2004. Predators select against high growth rates and risk-taking behaviour in domestic trout populations. Proceedings: Biological Sciences 271:2233.
Einum, S., and I. A. Fleming. 2000. Selection against late emergence and small offspring in Atlantic salmon (Salmo salar). Evolution 54:628–639.
Fleming, I. A., and M. R. Gross. 1994. Breeding success of hatchery and wild coho salmon (Oncorhynchus kisutch) in competition. Ecological Applications 3:230–245.
de Gaudemar, B., J. M. Bonzom, and E. Beall. 2000. Effects of courtship and relative mate size on sexual motivation in Atlantic salmon. Journal of Fish Biology 57:502–515.
Reisenbichler, R., S. Rubin, L. Wetzel, and S. Phelps. 2004. Natural selection after release from a hatchery leads to domestication in steelhead, Oncorhynchus mykiss. Pp. 371-383 in B. R. Howell, E. Moksness and T. Svåsand, eds. Stock enhancement and sea ranching. Oxford, Malden, MA.
Seamons, T. R., P. Bentzen, and T. P. Quinn. 2004. The mating system of steelhead, Oncorhynchus mykiss, inferred by molecular analysis of parents and progeny. Environmental Biology of Fishes 69: 333–344
Sundstrom, L. F., M. Lohmus, R. H. Devlin, and E. Brainerd. 2005. Selection On Increased Intrinsic Growth Rates In Coho Salmon, Oncorhynchus Kisutch. Evolution 59:1560.
Tymchuk, W. E., L. F. Sundstrom, and R. H. Devlin. 2007. Growth And Survival Trade-Offs And Outbreeding Depression In Rainbow Trout (Oncorhynchus Mykiss). Evolution 61:1225.
.Zimmerman, C. E., and G. H. Reeves. 2000. Population structure of sympatric anadromous and nonanadromous Oncorhynchusmykiss: evidence from spawning surveys and otolith microchemistry. Canadian Journal of Fisheries and Aquatic Sciences 57:2152–2162 | D | | Compare disease resistance | 07/01/2011 | 09/30/2011 | Concluded | Michael Blouin | 07/19/2010 12:21 PM |
| D | 162 | Analyze/Interpret Data | Data Analysis | Our data analysis efforts over the next year will focus on the following three actions:
(1) Estimate fitness of summer-run Hnew fish relative to that of wild summer-run
It is important to know whether our results with the winter-run steelhead are a special case, or are typical of hatchery supplementation programs. Several other research groups are in the process of examining the relative fitness of first-generation hatchery salmon of various species (coho, Chinook, steelhead), so eventually we will have a basis for comparison. But these studies have not published their results yet. We can now analyze the relative fitness of the first-generation summer-run steelhead stock that were released into the Hood River beginning in 1997. Those hatchery fish began returning in appreciable numbers in 2000. Because the winter and summer run are reproductively independent and breed in different parts of the Hood River, these summer-run data should constitute an independent test of the hypothesis that first-generation hatchery steelhead have lower fitness than wild fish. As part of the FY 2010 objectives we have been analyzing the first two run years (2000 and 2001) in which these summer-run Hnew bred in the wild (for these two years we should now have in hand >95% of their returning offspring). We will continue with the next year, with the ultimate goal of having four years of comparisons between Hnew summers and wild fish.
(2) Test whether offspring of hatchery fish grow faster under hatchery conditions than the offspring of wild fish
The 30-40% per generation drop in fitness that we estimated is occurring per hatchery generation in the winter run (Araki et al., 2007d) is so extreme that it was questionable whether domestication selection was even a plausible explanation. Here “domestication selection” means both positive selection for traits that are favored in the hatchery and relaxation of natural selection. We did a quantitative genetic analysis of selection in alternating environments and showed that strong selection was indeed a sufficient explanation for our results (Araki et al., 2008). So now the question becomes, what traits are involved?
Both adult traits and juvenile traits could be the targets of positive selection or relaxed natural selection. Artificial spawning in the hatchery almost certainly results in relaxation of mate selection (Berejikian et al.2000; de Gaudemar et al. 2000), intra-sexual competition, and selection on traits such as body size, egg size, fecundity and spawn timing and location (van den Bergheand Gross 1989; Fleming and Gross 1994; Einum and Fleming 2000). As for juvenile traits, given only 5-15% mortality in hatcheries, there is not enough selective death to generate the fitness declines observed. So the effect cannot be explained simply by viability (survival) selection on some trait in the hatchery. On the other hand, mortality from smolt to adult is often > 99%. So juvenile traits that affect adult survival (or perhaps even breeding behaviors) could well be the targets of selection. For example, smolt size is positively correlated with ocean survival, so there is strong selection on size at release (Reisenbichler et al., 2004). In the high-food and predator-free hatchery environment, this survival difference should select for high growth rate in hatchery juveniles, perhaps via behavioral or metabolic changes. But an excessively high growth rate can be maladaptive in natural environments (Arendt, 1997). So when those surviving hatchery fish reproduce in a natural setting, their offspring could have lower survival than wild offspring owing to inappropriate behaviors, an excessively high metabolism, or other maladaptive consequences of a genetic predisposition for fast growth in the benign and high-food hatchery environment (Sundstrom et al., 2005; Biro et al., 2004; Tymchuk et al., 2007). Therefore, we have begun testing whether there is evidence that the Hood River hatchery fish have been selected to have higher growth rates under hatchery conditions. In 2008 we set up a series of 2x2 matrix crosses involving a hatchery and wild male by a hatchery and wild female, and then raised each pooled group of four families in the hatchery for seven weeks. We measured the size of each fish and archived a tissue sample for DNA typing. After sorting them back into their respective families, we found no evidence for an additive genetic effect of hatchery versus wild background, although there was a weak maternal effect in which the offspring of wild females seemed to grow faster (perhaps an effect of egg size?). We started a second set of crosses in 2009 and sampled them at 2 months of age and one year of age. Among the 2-month olds we again found little evidence for a difference between fish of H vs. W backgrounds. We now have their siblings raised to one year of age, and plan to analyze them the same way. It is possible that any growth difference will appear later in life, so we haven’t rule out the growth rate hypothesis yet. Finally, we are setting up a 3rd set of crosses this summer, and will replicate the experiment in one final year, although raising them at densities that more closely match those of the production fish (last year’s fish were not as crowded). If we still see no effect of genetic background, then we will consider the growth rate hypothesis to be not so compelling.
(3) Test of differential disease resistance:
In collaboration with Jeri Bartholomew at OSU, we will attempt to test the siblings of this year’s crosses with viral and bacterial challenges. These experiments are to be conducted at the salmon disease lab at OSU. The fish will be challenged in groups and we will sort them back into their respective families after the fact via microsatellites, as with the growth rate study. Our hypothesis is that fish of H background will be less disease resistant as a consequence of hatchery selection for growth (i.e. selection to invest energetic resources into growth at the expense of maintenance). We haven't done this before, so we aren't guaranteeing that this first year's experiment will work (e.g. we have to estimate the proper challenge doses, fish can die for other reasons, and so on). But if it does work, and we do find a difference, it will be a very interesting corroborating evidence for selection on key life history traits as the mechanism of domestication.
References cited:
Araki, H., W.R. Ardren, E. Olsen, B. Cooper and M.S. Blouin. 2007a. Reproductive success of captive-bred steelhead trout in the wild: evaluation of three hatchery programs in the Hood River. Conservation Biology 21:181-190.
Araki, H., B. Cooper and M.S. Blouin. 2007d. Genetic effects of captive breeding cause a rapid, cumulative fitness decline in the wild. Science 318: 100-103.
Araki, H., B. Berejikian, M. Ford, and M.S. Blouin. 2008 Fitness of hatchery-reared salmonids in the wild. Evolutionary Applications 1:342-355.
Arendt, J.D.. 1997. Adaptive intrinsic growth rates: an integration across taxa. Quarterly Review of Biology 72:149-177.
Berejikian, B. A., E. P. Tezak, and A. L. LaRae. 2000. Female mate choice and spawning behavior of chinook salmon under experimental conditions. Journal of Fish Biology 57:647–661.
van den Berghe, E. P., and M. R. Gross. 1989. Natural selection resulting from female breeding competition in a Pacific salmon (Coho: Oncorhynchus kisutch). Evolution 43:125–140
Biro, P. A., M. V. Abrahams, J. R. Post, and E. A. Parkinson. 2004. Predators select against high growth rates and risk-taking behaviour in domestic trout populations. Proceedings: Biological Sciences 271:2233.
Einum, S., and I. A. Fleming. 2000. Selection against late emergence and small offspring in Atlantic salmon (Salmo salar). Evolution 54:628–639.
Fleming, I. A., and M. R. Gross. 1994. Breeding success of hatchery and wild coho salmon (Oncorhynchus kisutch) in competition. Ecological Applications 3:230–245.
de Gaudemar, B., J. M. Bonzom, and E. Beall. 2000. Effects of courtship and relative mate size on sexual motivation in Atlantic salmon. Journal of Fish Biology 57:502–515.
Reisenbichler, R., S. Rubin, L. Wetzel, and S. Phelps. 2004. Natural selection after release from a hatchery leads to domestication in steelhead, Oncorhynchus mykiss. Pp. 371-383 in B. R. Howell, E. Moksness and T. Svåsand, eds. Stock enhancement and sea ranching. Oxford, Malden, MA.
Seamons, T. R., P. Bentzen, and T. P. Quinn. 2004. The mating system of steelhead, Oncorhynchus mykiss, inferred by molecular analysis of parents and progeny. Environmental Biology of Fishes 69: 333–344
Sundstrom, L. F., M. Lohmus, R. H. Devlin, and E. Brainerd. 2005. Selection On Increased Intrinsic Growth Rates In Coho Salmon, Oncorhynchus Kisutch. Evolution 59:1560.
Tymchuk, W. E., L. F. Sundstrom, and R. H. Devlin. 2007. Growth And Survival Trade-Offs And Outbreeding Depression In Rainbow Trout (Oncorhynchus Mykiss). Evolution 61:1225.
.Zimmerman, C. E., and G. H. Reeves. 2000. Population structure of sympatric anadromous and nonanadromous Oncorhynchusmykiss: evidence from spawning surveys and otolith microchemistry. Canadian Journal of Fisheries and Aquatic Sciences 57:2152–2162 | E | DELIV | Complete interpretation of data | | 09/30/2011 | Concluded | Michael Blouin | 07/20/2010 11:15 AM |
| E | 161 | Disseminate Raw/Summary Data and Results | Communicate results | Post doc and/or principal investigator to attend at least one national or international scientific meeting for purpose of presenting current data and discussing work with colleagues. Meetings are usually in the summer months. It is important that the post doc and/or principal investigator meet regularly with colleagues who are doing similar work, and that we keep the scientific community informed about our latest results. This year we plan to attend at least one national or international meeting to to present our year-to-date results on the fitness of each type of fish. | A | | Attend meeting - info share | 10/01/2010 | 09/30/2011 | Concluded | Michael Blouin | 07/21/2010 9:44 AM |
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