| | | | | | | | | | | | |
|---|
| A | 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 | | 10/01/2009 | Concluded | Tracy Hauser | 06/29/2009 4:11 PM |
| B | 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 | 10/01/2009 | 10/01/2009 | Concluded | Tracy Hauser | 06/29/2009 4:11 PM |
| B | 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 | 10/01/2009 | 12/31/2009 | Concluded | Tracy Hauser | 06/30/2009 12:36 PM |
| B | 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 | 01/01/2010 | 03/31/2010 | Concluded | Tracy Hauser | 06/30/2009 12:36 PM |
| B | 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 | 04/01/2010 | 06/30/2010 | Concluded | Tracy Hauser | 06/30/2009 12:36 PM |
| B | 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 | 07/01/2010 | 09/30/2010 | Concluded | Tracy Hauser | 06/30/2009 12:36 PM |
| B | 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 | | 09/30/2010 | Concluded | Tracy Hauser | 06/29/2009 4:12 PM |
| C | 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 this year’s objectives we plan to analyze 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)
(2) Develop statistical methods for estimating hatchery contribution to resident parents
In the Hood River we find that even though we sample almost 100% of the anadromous fish, about 30% of the parents of returning adults are not in our sample of anadromous fish (see Table 4 in Araki et al. 2007a). Many more fathers are missing than mothers. These results suggest that the missing parents are resident (non-anadromous) fish. Other researchers doing pedigree work on steelhead find very similar patterns (e.g. Seamons et al., 2004; P. Moran, pers. comm.; E. Berntson, pers. comm.; D. Venditti, pers. comm.). Resident and anadromous forms in O. mykiss appear to be freely interbreeding life history forms in many populations of this species (Zimmerman & Reeves 2000). However, one important question for hatchery-supplemented rivers is whether those missing parents are naturally-produced trout or residualized hatchery fish that never went to sea. Therefore, one of our goals for the next year is to work on new statistical methods for using our multi-generation pedigree to estimate the proportion of those missing parents that were hatchery fish. If we are successful, we will estimate the proportion of missing parents that were hatchery fish.
(3) 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 preliminary experiments to test whether there is evidence that the Hood River hatchery fish have been selected to have higher growth rates under hatchery conditions. Last year 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. We now need to sort them back into their respective families via microsatellite genotyping in order to test if there is a main effect of fish type (hatchery, H, vs. wild, W) on growth rate. As part of this year’s work we propose analyzing these growth rate data.
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 | 07/01/2010 | 09/30/2010 | Concluded | Tracy Hauser | 06/29/2009 3:43 PM |
| C | 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 this year’s objectives we plan to analyze 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)
(2) Develop statistical methods for estimating hatchery contribution to resident parents
In the Hood River we find that even though we sample almost 100% of the anadromous fish, about 30% of the parents of returning adults are not in our sample of anadromous fish (see Table 4 in Araki et al. 2007a). Many more fathers are missing than mothers. These results suggest that the missing parents are resident (non-anadromous) fish. Other researchers doing pedigree work on steelhead find very similar patterns (e.g. Seamons et al., 2004; P. Moran, pers. comm.; E. Berntson, pers. comm.; D. Venditti, pers. comm.). Resident and anadromous forms in O. mykiss appear to be freely interbreeding life history forms in many populations of this species (Zimmerman & Reeves 2000). However, one important question for hatchery-supplemented rivers is whether those missing parents are naturally-produced trout or residualized hatchery fish that never went to sea. Therefore, one of our goals for the next year is to work on new statistical methods for using our multi-generation pedigree to estimate the proportion of those missing parents that were hatchery fish. If we are successful, we will estimate the proportion of missing parents that were hatchery fish.
(3) 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 preliminary experiments to test whether there is evidence that the Hood River hatchery fish have been selected to have higher growth rates under hatchery conditions. Last year 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. We now need to sort them back into their respective families via microsatellite genotyping in order to test if there is a main effect of fish type (hatchery, H, vs. wild, W) on growth rate. As part of this year’s work we propose analyzing these growth rate data.
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 | 07/01/2010 | 09/30/2010 | Concluded | Michael Blouin | 06/29/2009 4:45 PM |
| C | 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 this year’s objectives we plan to analyze 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)
(2) Develop statistical methods for estimating hatchery contribution to resident parents
In the Hood River we find that even though we sample almost 100% of the anadromous fish, about 30% of the parents of returning adults are not in our sample of anadromous fish (see Table 4 in Araki et al. 2007a). Many more fathers are missing than mothers. These results suggest that the missing parents are resident (non-anadromous) fish. Other researchers doing pedigree work on steelhead find very similar patterns (e.g. Seamons et al., 2004; P. Moran, pers. comm.; E. Berntson, pers. comm.; D. Venditti, pers. comm.). Resident and anadromous forms in O. mykiss appear to be freely interbreeding life history forms in many populations of this species (Zimmerman & Reeves 2000). However, one important question for hatchery-supplemented rivers is whether those missing parents are naturally-produced trout or residualized hatchery fish that never went to sea. Therefore, one of our goals for the next year is to work on new statistical methods for using our multi-generation pedigree to estimate the proportion of those missing parents that were hatchery fish. If we are successful, we will estimate the proportion of missing parents that were hatchery fish.
(3) 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 preliminary experiments to test whether there is evidence that the Hood River hatchery fish have been selected to have higher growth rates under hatchery conditions. Last year 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. We now need to sort them back into their respective families via microsatellite genotyping in order to test if there is a main effect of fish type (hatchery, H, vs. wild, W) on growth rate. As part of this year’s work we propose analyzing these growth rate data.
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 | | Develop statistical methods | 07/01/2010 | 09/30/2010 | Concluded | Michael Blouin | 06/29/2009 4:45 PM |
| C | 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 this year’s objectives we plan to analyze 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)
(2) Develop statistical methods for estimating hatchery contribution to resident parents
In the Hood River we find that even though we sample almost 100% of the anadromous fish, about 30% of the parents of returning adults are not in our sample of anadromous fish (see Table 4 in Araki et al. 2007a). Many more fathers are missing than mothers. These results suggest that the missing parents are resident (non-anadromous) fish. Other researchers doing pedigree work on steelhead find very similar patterns (e.g. Seamons et al., 2004; P. Moran, pers. comm.; E. Berntson, pers. comm.; D. Venditti, pers. comm.). Resident and anadromous forms in O. mykiss appear to be freely interbreeding life history forms in many populations of this species (Zimmerman & Reeves 2000). However, one important question for hatchery-supplemented rivers is whether those missing parents are naturally-produced trout or residualized hatchery fish that never went to sea. Therefore, one of our goals for the next year is to work on new statistical methods for using our multi-generation pedigree to estimate the proportion of those missing parents that were hatchery fish. If we are successful, we will estimate the proportion of missing parents that were hatchery fish.
(3) 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 preliminary experiments to test whether there is evidence that the Hood River hatchery fish have been selected to have higher growth rates under hatchery conditions. Last year 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. We now need to sort them back into their respective families via microsatellite genotyping in order to test if there is a main effect of fish type (hatchery, H, vs. wild, W) on growth rate. As part of this year’s work we propose analyzing these growth rate data.
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 growth | 07/01/2010 | 09/30/2010 | Concluded | Michael Blouin | 06/29/2009 4:45 PM |
| C | 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 this year’s objectives we plan to analyze 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)
(2) Develop statistical methods for estimating hatchery contribution to resident parents
In the Hood River we find that even though we sample almost 100% of the anadromous fish, about 30% of the parents of returning adults are not in our sample of anadromous fish (see Table 4 in Araki et al. 2007a). Many more fathers are missing than mothers. These results suggest that the missing parents are resident (non-anadromous) fish. Other researchers doing pedigree work on steelhead find very similar patterns (e.g. Seamons et al., 2004; P. Moran, pers. comm.; E. Berntson, pers. comm.; D. Venditti, pers. comm.). Resident and anadromous forms in O. mykiss appear to be freely interbreeding life history forms in many populations of this species (Zimmerman & Reeves 2000). However, one important question for hatchery-supplemented rivers is whether those missing parents are naturally-produced trout or residualized hatchery fish that never went to sea. Therefore, one of our goals for the next year is to work on new statistical methods for using our multi-generation pedigree to estimate the proportion of those missing parents that were hatchery fish. If we are successful, we will estimate the proportion of missing parents that were hatchery fish.
(3) 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 preliminary experiments to test whether there is evidence that the Hood River hatchery fish have been selected to have higher growth rates under hatchery conditions. Last year 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. We now need to sort them back into their respective families via microsatellite genotyping in order to test if there is a main effect of fish type (hatchery, H, vs. wild, W) on growth rate. As part of this year’s work we propose analyzing these growth rate data.
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 | | 09/30/2010 | Concluded | Tracy Hauser | 06/29/2009 3:43 PM |
| D | 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 | 05/01/2010 | 09/30/2010 | Concluded | Michael Blouin | 07/01/2009 1:53 PM |
| D | 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. | B | DELIV | Meeting attendance documented in final report | | 09/30/2010 | Concluded | Michael Blouin | 06/30/2009 5:43 PM |
| E | 119 | Manage and Administer Projects | Project/Contract Administration | Administrative work in support of on the ground efforts and in support of BPA's programmatic requirements such as metric reporting, financial reporting (e.g. accruals), and development of SOW package (includes draft and final SOW, budget and property inventory). | A | | Submit DRAFT 2011 SOW/budget to COTR | 07/01/2010 | 07/21/2010 | Concluded | Tracy Hauser | 06/29/2009 3:49 PM |
| E | 119 | Manage and Administer Projects | Project/Contract Administration | Administrative work in support of on the ground efforts and in support of BPA's programmatic requirements such as metric reporting, financial reporting (e.g. accruals), and development of SOW package (includes draft and final SOW, budget and property inventory). | B | FPCIR | Submit FINAL 2011 SOW/Budget to COTR | 07/21/2010 | 08/21/2010 | Concluded | Tracy Hauser | 06/29/2009 3:49 PM |
| E | 119 | Manage and Administer Projects | Project/Contract Administration | Administrative work in support of on the ground efforts and in support of BPA's programmatic requirements such as metric reporting, financial reporting (e.g. accruals), and development of SOW package (includes draft and final SOW, budget and property inventory). | C | ASSEB | Accrual - Submit FY 10 September estimates to BPA | 08/14/2010 | 09/10/2010 | Concluded | Tracy Hauser | 06/29/2009 3:49 PM |
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