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Pros vs. Cons of Hatchery Stocking: Part 2

8/14/2017

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PictureA nearly 18-inch stocked brook trout. What kind of havoc can these guys cause? Photo courtesy of PA Fish and Boat Commission.
Picking up where we left off last week, this week I’m going to flip the switch and talk about the potential negative consequences of stocking. I highly recommend that those who missed, or maybe just don’t remember, last week’s post start there before continuing. Some lingo and concepts may be a little fuzzy without the background information.
 
You didn’t press the back button, did you? (I don’t blame you, I wouldn’t either). So, here’s a quick refresher: wild trout populations have genes that are locally adapted to their native environments. This means that trout have genes that make them successful at life in their home stream, but their genes may not be great for surviving in another stream. However, not all fish in a stream can be identical clones of one another. There needs to be some level of genetic diversity in order for populations to survive disturbance and be able to adapt to future conditions. I call this the “eggs in many baskets” insurance policy. The genes that are best this year may not be the genes that are best next year, and so there needs to be high diversity so that at least some fish can survive and reproduce if conditions change in the future (if you’re a financial guru, this concept is very similar to having a diversified stock portfolio). 

Easy, right? Well, here’s some of the ways that hatcheries can disrupt this balance. I’m focusing specifically on recreational stocking programs because conservation stocking programs have taken more precautions to avoid these potential pitfalls (though, they do sometimes still happen).
  • Reduced genetic diversity: The goal of hatcheries is to grow as many big fish as possible. Once humans found out that growth was linked to genetics, we started weeding out the slow-growing fish, along with their genetics, leaving only a select few genes remaining in hatchery populations. This selection process (termed artificial selection) results in hatchery populations with desirable genetic make-ups, but very little genetic diversity. To think about this another way, imagine you have a bag of Skittles. If you scour through the bag and set aside every red Skittle, you’re left with a highly desirable handful of all-red candies. But, what if you want purple? Same idea with artificial selection in hatcheries.  The fish that are in our hatchery systems may have some highly desirable traits, but overall they have very little diversity and little ability to adapt to changing conditions. When these low-diversity hatchery populations are released into the wild, average population genetic diversity goes down.  
  • Introduction of maladapted genes: Not all genes are created equal.  Not only do we care about overall genetic diversity, but we also care about how much a particular gene increases survival and reproduction. Some genes decrease survival though, for example, making fish more predisposed to disease, heat stress, or predation. Many “bad” genes are either not present or in very low abundance in wild populations because natural selection removed them a long time ago. But, in hatcheries, there are very few selection pressures (aside from growth rate), and these genes have a tendency to pop up more frequently. Once hatchery fish are released into the wild, they bring these “bad” genes with them like an unwelcomed guest to the dinner party
  • ​Reduced survival and fitness of offspring: If hatchery fish never bred with wild fish, then the two aforementioned points would be of little concern. However, interbreeding between hatchery and wild fish is what causes those bad genes to enter into wild populations, and interbreeding is what causes the next generation of trout to have lower genetic diversity than offspring from two wild fish. I’ve made this analogy before, but think of offspring as an average of their two parents.  A fish that has two wild parents will receive mostly genes that have been naturally selected to increase survival in that stream.  Now think of a fish that has one wild and one hatchery parent. Half of that fish’s genes will be adapted for life in the stream, and the other half will be adapted for life in a concrete raceway. Because of this, fish that have one hatchery and one wild parent tend to have lower survival, lower reproduction, and, in general, less vitality, than fish that have two wild parents. This effect is known as outbreeding depression (breeding with fish OUT of the population depresses offspring survival), and the effects can be long-lasting. One generation of interbred fish with low survival will lead to fewer offspring for several more generations, and those future generations may still have lower genetic diversity and a higher rate of “bad” genes in their genetic composition.
Picture
Tiger trout are absolutely gorgeous, but I hate knowing that they steal from next year's native brook trout population.
  • Reduced fitness of wild populations: In addition to the genetic effects above, wild fish may invest a lot of energy into reproducing with hatchery fish, only to have few, if any, of their offspring survive. This energy, if spent reproducing with another wild fish, could have resulted in thousands of fertilized eggs with higher probability of survival. Sometimes we may not find many interbred individual in a population, and the initial reaction may be that hatchery fish are not negatively affecting wild populations.  But, it’s hard to tell how much a wild population would be reproducing if it wasn’t for their wasted efforts in trying to breed with hatchery fish. And, this potential negative effect extends beyond just brook trout stocking, but also to hybridization between brook and brown trout.  Every tiger trout (a brook x brown hybrid) represents a failed native brook trout reproduction. 
  • Decrease maximum size of wild trout:  This one might sound the most far-fetched, but hang with me. As I already mentioned, a trout’s maximum size has a genetic component.  Just like I am never going to be 6 feet tall, many trout will likely never reach 7 inches (the legal harvest size in most Pennsylvania streams). But, there are a few trout in every stream with the genetic potential to reach 7 inches and larger, and those fish may survive and reproduce to make more large fish for a very long time because most people don’t know they are there, and many wild trout anglers return catches of any size back to the stream.  Now, if we stock the stream, there is more angling pressure directed towards stocked fish, but the large wild trout are also more likely to get caught. And, because the wild trout are the size of hatchery trout, they are more likely get harvested because it’s assumed to be a hatchery fish. Eventually the select few fish that were carrying the “grow large” genes get removed from the population, and now no fish in the population has the genetic potential to get very large. . 
  • Increased competition: Alright, let’s move away from the genetic effects of hatcheries.  Hatchery fish are generally larger and more aggressive (which is, in part, the result of artificial selection). When they are released into the stream, they can outcompete native fish for food and occupy the best habitats in a stream. If hatchery fish remain in the stream for long, they can ultimately hurt native population reproduction by minimizing the amount of nutrients that wild fish can put towards growth and reproduction.
  • Predation: Bigger fish have a higher probability of being piscivorous, meaning fish-eating. Yes, wild fish eat juveniles, but the probability of this behavior occurring is much higher in stocked populations.
  • Disease: Incidences of disease are more common in hatchery populations, and stocking of diseased fish occurs fairly often.  In trout, the most common diseases tend to be gill lice and whirling disease, but there are also potential internal and fungal infections that don’t get as much attention due to their less-subtle appearance.   
I want to emphasize two points.  First, while these are the most likely negative effects of stocking, there are others that I didn’t cover. We would be here all day.  Second, not every stocking program is going to suffer from these negative effects. But, they are all possible and frequently observed. In the last part of this series I am going to attempt to pull together results from a few studies to comment on how likely it is hatchery stocking will have positive or negative effects on a wild populations.  But, I’m away at the annual meeting of the American Fisheries Society next week, so you may just have to wait in suspense….
 
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Pros vs. Cons of Hatchery Stocking: Part 1

8/5/2017

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PictureI caught this fish in Loyalsock in 2015 and learned this week that it's probably from a hatchery. Interesting, considering the stream isn't stocked.
Ask and you shall receive!  My recent posts have focused on the potential influence hatchery fish can have on wild populations, particularly if they starting reproducing with one another. With stocking a common, but often contentious, practice, I knew many people would be interested in our pending data analysis and results. But, I never thought it would get as much interest as it has. Even better, I’m getting some really great questions from many of you trying to wrap your head around the pros and cons of hatchery stocking.
 
Unfortunately, the manuscript I am in the process of writing will do very little to help clarify those questions. Heterozygosity, allelic richness, Hardy-Weinberg equilibrium, FST, genetic distance, bottlenecks …do these words mean much to you?  No?  That’s okay- they all refer to genetic measures that scientists look at to determine how an event (like stocking) is influencing a population. We know that certain events cause some of those numbers to go down, and other events will cause those numbers to go up.  Describing how those genetic statistics have changed (or not) as a result of stocking is largely what I’ll be talking about in my manuscript. Not very reader friendly.
 
But, those statistics are the nitty gritty. Just like you don’t need to know how an engine works to drive a car, you don’t really need to know the exact genetic details to understand the pros and cons of stocking. But, understanding some basic genetics concepts and a little lingo will go a long way in helping tease apart why the potential effects of stocking on native populations aren’t so cut and dry. It’s also helpful to have this background knowledge when deciding whether you feel the risks of stocking are worth the rewards. So, for the next three weeks I’m going to flesh out the possible pros (this week) and cons (next week), and then round out the mini-series by summarizing the consensus among biologists as to how much stocking is affecting fish populations worldwide.
 
Where to start. Let’s first talk a little about why we have to think about genetics when assessing the effects of stocking. The brook trout of today are the product of millions of years of natural selection- genes that produce healthier fish and the most offspring are more likely to get passed on to the next generation, whereas genes that are associated with lower survival and reproduction eventually get removed from the population. Making things more complicated, the best genes for one population are unlikely to be the best genes for another population. This is called local adaptation- millions of years of natural selection have left fish from a given stream with genes that give them the best chance of survival in that stream (highlighted for extra emphasis).  Local adaptation makes fish successful at living in one environment, but potentially not very successful if they are transferred into another environment. 

PictureAn example of the dangers of outbreeding depression- when individuals from different streams reproduce and the offspring have lower survival. Here, the yellow and red lines represent survival of the first- and second-generation offspring between two fish from different streams. You can see survival is dramatically lower than the original parents (blue and green lines). Image from conservationbytes.com.
Following that line of thought, it seems like fish should stay in their home stream to fine-tune local adaptation and increase survival.  But, it’s not that simple (for starters, fish don’t choose to adapt, but that’s a story for another day). Streams are highly variable environments, and local conditions change faster than fish can adapt. So, stream fish populations need to put their eggs in more than one basket- they need some fish that have genes that are successful under certain conditions and other fish that are successful in other conditions. In other words, populations need a lot of genetic diversity. With increased genetic diversity there is increased survival and resiliency to changing conditions, but also increased potential for populations to be able to adapt to future stressors (like climate change).
 
A lot of things influence genetic diversity, but some of the biggest contributors are population size and population connectivity. Big populations that have a lot of fish moving into and out of them tend to have high genetic diversity.  Historically, this is probably how many brook trout populations existed.  But, times have changed. Population sizes have declined following natural disaster, disease, habitat loss, etc. And, many populations are now isolated by waterfalls, road crossings, and thermally unsuitable habitat. When populations get too low and too isolated, genetic diversity quickly erodes as fish start inbreeding and other genes get randomly removed from the population (a process known as genetic drift).  For all of these reasons, one of the biggest priorities fisheries conservation managers have is to restore genetic diversity by increasing population size and connectivity. 
 
Did you follow all that?  No?  That’s okay. Big picture- fish become genetically specialized to the local environment (local adaptation), which increases their survival. But, the population can’t become too specialized because it needs high genetic diversity in order to be withstand disturbance and have the potential to adapt to future conditions.
 
Okay, so where does stocking come into play?
 
First off, there are two basic forms of fish stocking. It’s important to keep them separated as we discuss the pros and cons of stocking because they are very different from one another and have their own benefits and drawbacks. The less common form is conservation stocking, where the goal is to increase population sizes and genetic diversity of critically threatened or endangered fish or reintroduce a species to its native habitat after restoration. In conservation stocking, the fish populations are generally not harvested, and stocking is used to prevent future population declines and extinction.
 
Conservation stocking is very tricky business. The genetics of every individual used for reproduction are carefully considered so that the stocked population is locally adapted to the wild environment, thus giving fish the best chance of surviving and reproducing. This also helps avoid outbreeding depression - when a native, locally adapted fish spawns with stocked fish that is not locally adapted and the offspring have lower survival and reproduction. Outbreeding depression has the potential to cause rapid population declines as each generation continues to have lower and lower survival and reproduction. To avoid potential effects of outbreeding depression, fish used in conservation stocking are often brought in directly from the wild and only kept in captivity for a few generations, thus minimizing genetic differences between wild and stocked populations. 

PictureYellowfin madtom is just one example of a species being stocked for conservation. Recently populations were released into their native range in Virginia, and the success of the project is still being monitored. Photo from knoxmercury.com.
Now, contrast that with recreational stocking. Recreational stocking, especially for trout, is by far the most common. The goal here is to stock as many large fish as possible in order to increase angler satisfaction. Genetics are considered, but mostly as a means to grow bigger fish, faster. In other words, fish used for recreational stocking have been artificially selected.  As opposed to natural selection (where the environment picks the best genes), in artificial selection humans are the ones deciding which genes are best. It takes many years for humans to artificially select the genes that will make a population grow large and fast. But, once that goal is achieved, wild fish are no longer brought into the hatchery because wild fish will not grow as large and as fast as their artificially selected counterparts. In fact, it’s been over 100 years since the last wild fish has been introduced into many hatcheries used for recreational stocking. Because of this, fish used in recreationally stocking are genetically incompatible with the wild environment, and have lower survival and reproduction once released.
 
So, what are the pros of stocking?
  1. Increased recreational opportunities. This is an obvious one, but it’s an important one. One of the reasons I find trout management so fun (and sometimes so frustrating) is that it’s a fish prized by anglers.  It is important to offer people the opportunity to catch brook trout so we can carry on traditions and provide opportunities for people to enjoy nature, and sometimes that’s most feasible with stocking.
  2. Increased local population sizes: Similar to the point above, stocking does increase local population sizes. This may be temporary as stocked fish are harvested or die shortly after stocking, but the increased population size is sometimes necessary to allow streams to be fished. As brook trout populations continue to decline, many streams would probably be closed for angling if it were not for the ability to supplement wild populations with stocked. And, if stocked fish survive, then the increased population size can decrease the chances of inbreeding (but come back next week to learn why this could be bad).
  3. Increase genetic variability: If stocked fish have more genetic variability than the native population, then stocking can increase populations genetic diversity which, in turn, can increase overall population health. For the reasons described above, this is usually not the outcome of stocking.  But, again, I’ll discuss this more next week.
  4. Increased resistance to disease or stress: While artificial selection does not usually increase survival of fish in the wild, it is possible to artificially breed fish that are resistant to many diseases that can devastate wild populations. Likewise, we can sometimes breed fish that are more tolerant to environmental stressors, such as heat. These stocking goals are just starting to become possible, but could be more realistic in the future when we know more about specific genes that lead to desired characteristics. But, spoiler alert, there are limitations to this practice. For example, I don’t think we’ll ever produce a brook trout that can survive in warmwater streams.
  5. Populations recovery: This one focuses specifically on conservation fisheries. Many populations of threatened and endangered species have been improved through stocking efforts. And, in many cases, this recovery probably would not have been possible without the assistance of stocking.
 
So, yes, stocking can have positive influences on native populations. But, I’ve already hinted a few times that most stocking and hatchery practices are unable to realize some of these potential benefits. But, tune in next week for a more detailed discussion about the possible negative consequences of stocking. 


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