Anemone Reproduction

MarineBioHSU

In Memoriam
Ok folks, so I am starting this thread with the hopes that we can as a community try and uncover and quantify what factors initiate reproduction in anemones in our aquariums, both sexual and asexual. Obviously, a good place to start is to understand what factors initiate reproduction events in the wild, and try and correlate this to captivity.

I am currently involved in farming E. quadricolor via manual division, and while I think that almost any species of anemone can undergo transvers fission and survive, I do not feel that at this point, it is the best method of farming these guys in captivity. Instead, I feel we should strive to lean how to consistently stimulate spawning in captivity. There are numerous accounts of captive spawning occuring, albeit unintended by most aquarists, however it is an encouraging sign that it is possible. Additionally, Anna Scott has done a fair amount of research into sexual reproduction of several host anemone species in captiviy, apparently with a fair amount if success.

So, it seems like one of the primary points of interest/contention is what exactly causes anemones (namely E. quadricolor) to reproduce via transverse fission.

There are numerous reports of this happening in captivity, and many of the reports seem contradictory.

One of the general ideas posed by biology is that, in general, a species will reproduce via asexual means when conditions are suitable enough that no exchange of gametes are necessary. Where-as sexual reproduction would then be stimulated by a stressful event that would necessitate a mixing of genetic material, either to deal with a reduced genetic pool, or perhaps to produce encysted offspring that can withstand a period of toxicity.

At first glance this would seem to indicate that BTA's for example, would be dividing in our tanks as a result of favorable conditions, such as enough food etc. It does seem to make sense that the anemones could be "thinking" that if their genotype is suitable to their environment, there is no reason to alter things. One piece of evidence that seems to support this are the numerous reports from aquarists that feeding their anemones heavily for a week or so will result in a splitting event.

HOWEVER, at the very same time many aquarists feel that their anemones split as a result of stress, for example some folks notice a large water change, power outage etc can stimulate a division. Furthermore, some aquarists feel that "overfeeding" can stress anemones into dividing, which seems like it may not make a lot of sense. However, again it has been so well documented in captivity that there is clearly some merit.

And perhaps most interesting of all, many aquarists that are successful with anemones have very different experiences, what I mean is some times a well fed BTA may continue to grow and grow and never divide, where-as another well fed and happy anemone will divide like no other! To me this would indicate that there are other factors coming into play.

So, in light of the above evidence I would like to examine what benefits and drawbacks there are to different methods of reproduction for anemones.

1. Asexual reproduction: By and large all the anemones that we keep in captivity that can/do reproduce asexually undergo transverse fission, that is they divide through the mouth, however there are reports of budding and intratentacle polpys being formed as well.

Again, standard biological wisdom would seem to indicate this method of reproduction would occur during times of plenty. However several studies have indicated there could be other factors. For example, perhaps the anemones divide as a means of tissue regeneration, which theoretically could explain division from "overfeeding"....maybe after digesting food the anemone "feels" a need to re-grow GVC tissues that were utilized, and division is a good method to speed this up?

Furthermore, several studies have shown a positive inverse correlation between feeding and division rates, that is to say, in the experiements, anemones that were not fed divided frequently, where-as anemones that were fed often did not divide. This would seem to contradict many of the observations from hobbyists, myself included.

An additional study done on the east coast of North America showed that anemones in higher water flow divided more rapidly thatn those found in lower flow situations, and this would seem to refelct the conventional idea held that smaller colonial BTA's are found in shallow waters and larger individual specimens tend to occur in deeper waters.

So to try and sum up asexual division:

1. Some evidence suggests asexual division may occur as a result of preserving genotypes that are suited to current conditions.

The idea behind this theory is that if the anemones are doing well in their environment, then there is no reason to change so to speak, in other words if it aint broke dont fix it. This could explain why anemones in shallow water/high flow situations divide more often, more food being swept into their tentacles/more light etc. This would of course explain the observations that increased feeding induces splitting in some captive anemones.

2. Some evidence suggests that asexual division may occur as a means of growth/tissue regeneration.

This theory proposes that old tissue can be shed via division, the net result being two individuals with a significant amount of new tissue/growth. Since anemones have no know life span and indeterminate growth, this theory makes sense.

This theory may be harder to test in captivity, but not impossible. I propose that anemones under higher lighting situations may need to divide more frequently for tissue regeneration as a means of dealing with free radicals resulting from UV light/photosynthesis etc.


3. Some evidence suggests that asexual divison may occur as a last ditch effort to preserve an individuals genotype when faced with degrading conditions.

This theory could explain why anemones may divide under stressful situations. Say for example that conditions are rapidly becoming unfavorable, and the anemone does not have time to release gametes, its best option would be to try and divide rapidly to give its individual genotype the best chance of survival.

There are a few problems I see with this theory:
1. Many studies have shown that most anemones almost constantly have mature gametes present in their mesentaries.
2. Additionally, a spawning event would produce a far greater amount of offspring, therefore a greater chance of survival, in addition to a greater chance that a beneficial mutation could occur that would create a genotype more suited for altering conditions.
3. Numerous observations of known stressful occurences (such as el nino years) on reefs have shown that when faced with such conditions, host anemones tend to expell zoanthellae and maintain themselves via feeding and self absorbtion. So, it would seem like reducing their nutrient reserves through division and the energy required to divide and heal would not be a "wise choice" in unfavorable conditions.

However, there are a few possible explainations for this idea:
1. Some studies have shown that the smaller the diameter of an anemone, the more effective the gain from food capture. So maybe if resources are diminishing, then by dividing, multiple smaller anemones would be able to sustain themselves longer on a single morsel of food than a single large anemone could on the same resources.


At this point it is certainly impossible to make any sort of absolute conclusion, however in light of the evidence, I would suggest that we can start to piece together different scenarios leading to division.

For example, it seems to me that BTA's kept in high light, high flow situations and fed often will divide frequently. Where-as anemones kept in lower light, lower flow situations and still fed frequently may opt to grow instead of divide. Many possibilities.

So, now I am asking everyone here to contribute their ideas/observations. I would also like to come up with a set of parameters to watch/examine that may cause division, or not.

For example, large water changes seem to stimulate division. Perhaps this could be a result of reducing the chemotoxins in the water, thus giving the anemones the impression that there is more room/less competition so they can divide?

Or perhaps a stressful situation such as being left in a bucket or a power outage could cause tissue necrosis, and the anemones response could be to divide, thus ridding itself of the dying tissue and jump starting an immunological response? Again, just my ideas.

This afternoon I will post a list of books/articles/journals etc that I have been reading so that y'all can look for them as well.

I look forward to any and all idea, observations and opinions. I think the knowledge base here is extraordinary and if we can compile and share these ideas in an organzied manner, we can make a lot of progress.

Thanks.
 
MarineBioHSU;18660274 Additionally said:
Studies? what does her research tell us about sexual reproduction in captive host anemones?[/COLOR]


So, it seems like one of the primary points of interest/contention is what exactly causes anemones (namely E. quadricolor) to reproduce via transverse fission.

More important in my mind.. What causes them to undergo sexual reproduction?

There are numerous reports of this happening in captivity, and many of the reports seem contradictory.

One of the general ideas posed by biology is that, in general, a species will reproduce via asexual means when conditions are suitable enough that no exchange of gametes are necessary. Where-as sexual reproduction would then be stimulated by a stressful event that would necessitate a mixing of genetic material, either to deal with a reduced genetic pool, or perhaps to produce encysted offspring that can withstand a period of toxicity.

Using general biology principles to try and explain sexual reproduction in anemones may not be all that applicable as they dont always "follow the rules"..

At first glance this would seem to indicate that BTA's for example, would be dividing in our tanks as a result of favorable conditions, such as enough food etc. It does seem to make sense that the anemones could be "thinking" that if their genotype is suitable to their environment, there is no reason to alter things. One piece of evidence that seems to support this are the numerous reports from aquarists that feeding their anemones heavily for a week or so will result in a splitting event.

HOWEVER, at the very same time many aquarists feel that their anemones split as a result of stress, for example some folks notice a large water change, power outage etc can stimulate a division. Furthermore, some aquarists feel that "overfeeding" can stress anemones into dividing, which seems like it may not make a lot of sense. However, again it has been so well documented in captivity that there is clearly some merit.

Here is how I see it with BTA's: They seem to prefer asexual reproduction to sexual reproduction.. All anemone species dont share the same characteristics and preferences for their reproduction. I have never heard of BTA's sexually reproducing in captivity (im sure it has happend). BTA's tend to split, both when they are doing well as a means of growing their population, and they also seem to split under stress for increased survival.



And perhaps most interesting of all, many aquarists that are successful with anemones have very different experiences, what I mean is some times a well fed BTA may continue to grow and grow and never divide, where-as another well fed and happy anemone will divide like no other! To me this would indicate that there are other factors coming into play.

I dont see these hobbyist experiences as contridictory. I dont think we can conclude a modal preference of BTA reproduction from this.

So, in light of the above evidence I would like to examine what benefits and drawbacks there are to different methods of reproduction for anemones.

I assume you are talking about only BTA's?

1. Asexual reproduction: By and large all the anemones that we keep in captivity that can/do reproduce asexually undergo transverse fission, that is they divide through the mouth, however there are reports of budding and intratentacle polpys being formed as well.

Again, standard biological wisdom would seem to indicate this method of reproduction would occur during times of plenty.


However several studies have indicated there could be other factors. For example, perhaps the anemones divide as a means of tissue regeneration, which theoretically could explain division from "overfeeding"....maybe after digesting food the anemone "feels" a need to re-grow GVC tissues that were utilized, and division is a good method to speed this up?

Furthermore, several studies have shown a positive inverse correlation between feeding and division rates, that is to say, in the experiements, anemones that were not fed divided frequently, where-as anemones that were fed often did not divide. This would seem to contradict many of the observations from hobbyists, myself included.

(sources?) It is fine for you to copy and past one/several paragraphs from your sources that illustrate the point you are trying to make.. (not illegal i mean). This way we can read along with what you are trying to say from the literature that says it.

An additional study done on the east coast of North America showed that anemones in higher water flow divided more rapidly thatn those found in lower flow situations, and this would seem to refelct the conventional idea held that smaller colonial BTA's are found in shallow waters and larger individual specimens tend to occur in deeper waters.

I see this as a SA/V deal.. more environmentally active conditions
(high flow) might favor a larger SA/V ratio for gelatinous creatures like anemones (just a thought).

So to try and sum up asexual division:

1. Some evidence suggests asexual division may occur as a result of preserving genotypes that are suited to current conditions.

The idea behind this theory is that if the anemones are doing well in their environment, then there is no reason to change so to speak, in other words if it aint broke dont fix it. This could explain why anemones in shallow water/high flow situations divide more often, more food being swept into their tentacles/more light etc. This would of course explain the observations that increased feeding induces splitting in some captive anemones.

2. Some evidence suggests that asexual division may occur as a means of growth/tissue regeneration.

This theory proposes that old tissue can be shed via division, the net result being two individuals with a significant amount of new tissue/growth. Since anemones have no know life span and indeterminate growth, this theory makes sense.

This theory may be harder to test in captivity, but not impossible. I propose that anemones under higher lighting situations may need to divide more frequently for tissue regeneration as a means of dealing with free radicals resulting from UV light/photosynthesis etc.


3. Some evidence suggests that asexual divison may occur as a last ditch effort to preserve an individuals genotype when faced with degrading conditions.

This theory could explain why anemones may divide under stressful situations. Say for example that conditions are rapidly becoming unfavorable, and the anemone does not have time to release gametes, its best option would be to try and divide rapidly to give its individual genotype the best chance of survival.

There are a few problems I see with this theory:
1. Many studies have shown that most anemones almost constantly have mature gametes present in their mesentaries.
2. Additionally, a spawning event would produce a far greater amount of offspring, therefore a greater chance of survival, in addition to a greater chance that a beneficial mutation could occur that would create a genotype more suited for altering conditions.
3. Numerous observations of known stressful occurences (such as el nino years) on reefs have shown that when faced with such conditions, host anemones tend to expell zoanthellae and maintain themselves via feeding and self absorbtion. So, it would seem like reducing their nutrient reserves through division and the energy required to divide and heal would not be a "wise choice" in unfavorable conditions.

However, there are a few possible explainations for this idea:
1. Some studies have shown that the smaller the diameter of an anemone, the more effective the gain from food capture. So maybe if resources are diminishing, then by dividing, multiple smaller anemones would be able to sustain themselves longer on a single morsel of food than a single large anemone could on the same resources.

I like these theories and dont see any of them as mutally exclusive

At this point it is certainly impossible to make any sort of absolute conclusion, however in light of the evidence, I would suggest that we can start to piece together different scenarios leading to division.

For example, it seems to me that BTA's kept in high light, high flow situations and fed often will divide frequently. Where-as anemones kept in lower light, lower flow situations and still fed frequently may opt to grow instead of divide. Many possibilities.

IME, with bublle tips anemones I have seen this often as well

So, now I am asking everyone here to contribute their ideas/observations. I would also like to come up with a set of parameters to watch/examine that may cause division, or not.

For example, large water changes seem to stimulate division. Perhaps this could be a result of reducing the chemotoxins in the water, thus giving the anemones the impression that there is more room/less competition so they can divide?

Or perhaps a stressful situation such as being left in a bucket or a power outage could cause tissue necrosis, and the anemones response could be to divide, thus ridding itself of the dying tissue and jump starting an immunological response? Again, just my ideas.

This afternoon I will post a list of books/articles/journals etc that I have been reading so that y'all can look for them as well.

I look forward to any and all idea, observations and opinions. I think the knowledge base here is extraordinary and if we can compile and share these ideas in an organzied manner, we can make a lot of progress.

Thanks.
 
Don't overlook that fact that at least one very well respected anemone biologist has stated that she has a suspicion that E. quadricolor might/should be divided into two species. A smaller species that divides readily and a large species that doesn't and possibly can't divide as a means of reproduction.

Looking for causes of spontaneous fission may be futile. One explanation as to why we see different causes for BTAs to divide is that there is no initiating factor. They may divide because that is just how they grow. BTA's that don't divide when presented with the same factors may well be the version of BTA that doesn't/is not able to divide.

I am not saying you should stop your study. You do need to keep your mind open to other possibilities though. You already have a lot of data. I am not sure you are going to get anything different from the board member than what you already have. If you have a large supply of BTAs at your disposal, it may be time for you to design a controlled experiment. Can you say Master's Theseus?
 
and while I think that almost any species of anemone can undergo transvers fission and survive

You stopped me in my tracks right here :) Before I go on and read further, let me guarantee you this is NOT the case. You have experience with one of two clown anemones that asexually reproduce with frequency in the wild. Just because you can cut an E. quadricolor in half and reasonable assume both halves will survive does NOT mean that the said can be said for other species.
 
Ok, having read your entire note here are my comments:

(1) I like the logical approach you are taking to try to understand what is witnessed frequently, but not understood.

(2) Are you interested in the theoretical, or the practical application of this knowledge?

(a) If theoretical, then go for it.

(b) If practical, why does it matter? If E. quadricolor can be manually divided with a high success rate, why is it important to trigger the event naturally? To what benefit?

(3) Why are you focusing on asexual versus sexual reproduction?
 
Ok, having read your entire note here are my comments:
(2) Are you interested in the theoretical, or the practical application of this knowledge?
(a) If theoretical, then go for it.

Agreed with this and the next point.


Ok, having read your entire note here are my comments:]
(b) If practical, why does it matter? If E. quadricolor can be manually divided with a high success rate, why is it important to trigger the event naturally? To what benefit?

Also agreed. We have the knife/scissor method down with very high success rates. Stimulating natural asexual reproduction, for practical application, is not necessary, IMO. But, if you are interested in the theory behind it and trying to "unlock the secrets" (as mentioned above) then, yes, go for it.


(3) Why are you focusing on asexual versus sexual reproduction?

Right. It seems you have started off wanting to address and figure out sexual reproduction but then went to trying to unlock asexual reproduction.

While both methods of reproducing, most likely many of the same events that trigger spawning in corals - long-term seasonal fluctuations, lunar cycles, sunsets/sunrises, chemical messaging, colony/individual size and nutrition, etc. - will also trigger spawning in anemones. I don't know that the same can be said for asexual reproduction - though I don't know for sure, but I doubt it.

In addition - perhaps to our advantage, stress spawns will also occur. This seems to be what happens most of the time in captivity and needs to be taken advantage of whenever possible - especially since mimicking natural reef conditions are nearly impossible.

S. haddoni seems to be the best candidate for the host anemones.

Cheers
Mike
 
By the way - if my post sounded harsh, it wasn't intended as such. I think this is a good goal and toward the direction for compiling information. I was just trying to understand some of the thought process.

Cheers
Mike
 
Thanks for all the input guys.

Phender: Good point, however I do not feel there is enough data to make a definitive dtermination either way. My understanding is that the idea is based solely on size/morphological differences between shallow and deep water specimens. And again, there are studies showing that anemones moved from deeper/slower flow areas to shallower/faster flow areas will begin to divide more often and vice versa.

I don't think looking fro the cause of asexual fission is futile, I think it is very doable and indeed crucial to a better unerstanding of these guys. I think there may be a variety of situations that lead to asexual fission in captivity, and I would like to try and put the pieces together. I just don't want to hear folks telling me and others that my anemones must not be well taken care of if they are dividing, because they are constantly stressed.

BonsaiNut: What I mean is that there is significant evidence that suggests that almost if not all anemones are capable of dividing asexually on their own, in the wild. I base this on what I have read and from what I have been told from folks with first hand observations. I do not think we should attempt to divide them in captivity until we know a lot more about their immunological responses etc. I am sick and tired of hearing people say "hey look I cut my carpet in half yesterday and it looks great, I just sliced it and threw it back in my display tank!", of course they invariably die in the long run, I think however it may be valuable to undertake such a study in a well contolled experiment done with proper facilities etc. Atleast that way real data can be obtained.

I am of course interested in both theoretical and applicable data. I am a biologist, if I see a fish eat its own poo I wonder why lol. In this case I think all knowledge gained here is applicable. Yes, BTA's can be divided manually with good success, but this has long term implications. Also, BTA's are a great candidate for working with triggering sexual reproduction in captivity due to their relative ease of maintainance.

I am focusing on asexual repro. here due to the sheer number of times it has happened in captivity, vs. sexual reproduction, which ahs occured but is much less documented or understood. Not saying that I don't have ridiculously expensive controllers on some dedicated systems trying to initiate a spawning event lol.

ousnakebyte: Not harsh at all! Again I appreciate any and all input.

Thanks again.
 
Thanks for all the input guys.

............

I don't think looking fro the cause of asexual fission is futile, I think it is very doable and indeed crucial to a better unerstanding of these guys. I think there may be a variety of situations that lead to asexual fission in captivity, and I would like to try and put the pieces together. I just don't want to hear folks telling me and others that my anemones must not be well taken care of if they are dividing, because they are constantly stressed.

................

I didn't say it is futile. I said it may be futile if, as people's experience suggests (at least to me), there doesn't seem to be a consistent trigger, but instead is just part of their growth pattern.
Maybe colonial BTA's act more like corals in their growth than solitary anemones. I don't know why corals form colonies of polyps instead of one polyp getting bigger and bigger, but maybe colonial BTAs have the same survival adaptation.

I am thinking that the only way for you to get an answer is to do a controlled experiment. However, most experimental results/conclusions I have read simply lead to more questions and few conclusions. I am just trying to throw stuff at you that you may not have considered, so when you actually do the experiment, you will get some meaningful results and not just questions that others have already asked.

Good luck! I have a BTA that I (and others) would love to have split. It hasn't split in the 5 years I have had it and I am not about to cut it, just in case it happens to be one of the non-splitting BTAs(if there is such a thing). If you could figure out the trigger, you would make a lot of people on my waiting list very happy. :)
 
Phender: Good point, however I do not feel there is enough data to make a definitive dtermination either way.

This, I totally agree with. As of right now, it's just a matter of personal opinion, or what one wants to believe.

My understanding is that the idea is based solely on size/morphological differences between shallow and deep water specimens.

And their method of reproduction.

And again, there are studies showing that anemones moved from deeper/slower flow areas to shallower/faster flow areas will begin to divide more often and vice versa.


Please post links to the studies describing this. There are several species that demonstrate the same characteristics of having smaller individuals that reproduce asexually on a regular basis, and larger ones that don't. E. quadracolor, H. magnifica, and tapetum, just to name a few. I'm sure there are others. I've never read a study describing the behavior you're referring to.

I don't think looking fro the cause of asexual fission is futile, I think it is very doable and indeed crucial to a better understanding of these guys. I think there may be a variety of situations that lead to asexual fission in captivity, and I would like to try and put the pieces together.

I wish more hobbyists would take the time to do what you're attempting. The hobby would probably be much further along by now if there were.:thumbsup:

I just don't want to hear folks telling me and others that my anemones must not be well taken care of if they are dividing, because they are constantly stressed.

How could anyone make such statements without knowing the details of the particular system and husbandry? I simply wouldn't listen to people that would make such statements.

BonsaiNut: What I mean is that there is significant evidence that suggests that almost if not all anemones are capable of dividing asexually on their own, in the wild.

If there is "significant evidence that suggests that almost if not all anemones are capable of dividing asexually on their own, in the wild", you should have no problem providing the links that would support your statement. I have been reading research papers on this subject for years, and have never ran across one that would support this notion.
 
This subject is so huge it is hard to summarize :) Additionally, though there is some material available, it often deals with temperate water anemones. Many/most of these anemones do not harbor zooxanthellae, which may have a significant impact on reproductive behavior. There is little available for clown anemones that I am aware of. Here are some additional thoughts.

(1) Reproduction among anemones varies substantially within a genus, and may even vary within species. Some anemones exhibit different types of reproduction including more than one method of asexual reproduction.

(2) There are substantial energy costs associated with reproduction. Vegetation reproduction (i.e. cloning) has a lower energy cost than sexual reproduction. However there are costs and benefits to each method - the primary benefit of sexual reproduction being genetic diversity.

(3) Anemone species that rely wholly on sexual reproduction have larger oocytes (eggs) than those that combine sexual and asexual reproduction. Anemones with small oocytes are more likely to be brooders than those with large oocytes. Very few anemone species rely solely on asexual reproduction.

(4) Asexual reproduction is often triggered by stress - including stress associated with lack of food. However it can also be triggered by an abundance of food. Different anemones prioritize growth over asexual reproduction (and vice versa) when presented with high food availability.

(5) For anemones that reproduce sexually and asexually, asexual reproduction is correlated with small size. The more optimal the environmental conditions, and the larger the individual, the more likely it will reproduce sexually versus asexually. This has led some to surmise that large individuals may be a different species/sub-species, but genetic research has proven this to not be the case (at least not always).

(6) Because of the relationship between size, energy cost of living, energy availability, and energy cost of reproduction, there appears to be a formula dictating the optimal and maximum size of anemones based on their species and environment (Sebens, 1979). In times of stress, asexual reproduction can be an "emergency" mechanism to reduce body mass and ensure survival of species by creating numerous low energy individuals instead of one large high energy individual.

(7) Some anemones can reproduce continually (sexually and/or asexually) while others do so seasonally. Higher temperatures (within a fixed feeding regimen) tend to increase the rate of asexual reproduction, while lower temperatures tend to lower it.

(8) When presented with high food availability, some anemones prioritize growing larger versus asexually reproducing (which lowers body size). This may be due to seasonal or other variations where in times of plenty the anemone reproduces sexually, and in times of famine the anemone reproduces asexually and lowers body size to become more efficient.

There is so much out there on this subject that it is like peeling an onion. I have read probably 20 or 30 papers so far and there is more out there. Unfortunately because of the amazing diversity of anemones, and the lack of specific work on clown anemones, you are left wondering which research applies to tropical clown anemones.

Frankly, I refer to Anna Scott's work on sexual reproduction simply because it deals definitely with H. crispa and E. quadricolor. E. quadricolor is a prolific asexual reproducer, while H. crispa is not known to reproduce asexually.
 
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Hey folks, thanks for all the replies. I am so sorry I have not responded yet, I have been busy as heck at school and sick as a dog, too much time packed in the library with other sick people studying.

At any rate, I am gathering all the references etc to post them for you, and just wanted to let y'all know I really appreciate the interaction and advice from all you experienced nem owners! I will hopefully be able to post said links this evening.

Thanks for your patience.
 
HOWEVER, at the very same time many aquarists feel that their anemones split as a result of stress, for example some folks notice a large water change, power outage etc can stimulate a division. Furthermore, some aquarists feel that "overfeeding" can stress anemones into dividing, which seems like it may not make a lot of sense. However, again it has been so well documented in captivity that there is clearly some merit.


Thanks.

Last year I used to own a number of E quadricolor (Cloned from another reefer), and I did feed heavily (everyday with krill and mysis) and they did not split. I also performed 10-15 gallon water changes every week on the 60 gallon system and they did not split.
 
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