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Invisibletexas34
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Bifactorial mating system
    #5599240 - 05/06/06 10:32 PM (18 years, 16 days ago)

Hi, I'm a longtime reader of the boards that finally registered a few months back and only recently started posting. I've always been impressed with the site, its members, and the discussions taking place. There are a great many people here who know more than I do about many things; however, there are some subjects to which I can add meaningful information. One of these subjects is bifactorial incompatibility aka tetrapolar incompatibility, the mating type system found in most mushrooms.

Two fairly recent papers from the James lab at Duke give an excellent summary of this mating system in their introductions. The intros are complementary and give a nice overview together. These papers are freely available as .pdfs Here and Here. Because the authors write a better explanation that I would, here is a short quote from the fist link,
Quote:

The homobasidiomycetes,or mushroom fungi, have arguably the most complex mating system of all known organisms. Many species possess a mating system known as bifactorial (or tetrapolar) incompatibility, where two unlinked loci control the mating-type of an individual. Compatibility between mates occurs only when they differ in alleles at both of the incompatibility loci (the A and B mating-type loci). Mushroom incompatibility loci are also distinctive in that many species have over 100 mating-type alleles per incompatibility locus (Raper, 1966).



And one from the second link,
Quote:

In large part, this variation can be explained by the structure of the mating-type loci in which each mating-type locus is comprised of two or more closely linked subloci (Halsall et al., 2000; K€ues and Casselton, 1993). Recombination between subloci creates new combinations of alleles at the subloci; each combination confers a unique mating-type specificity. As a result of the high number of mating-types, any two haploid individuals that encounter each other in nature are likely to be compatible. The requirement for heterozygosity at the mating-type loci allows the maintenance of numerous mating-types due to balancing selection. The action of balancing selection extends the life of each mating-type allele so that its duration in the population is over a much longer time scale than an allele at a neutral locus (May et al., 1999). This, in turn, causes a large level of sequence divergence between mating-types due to their ancient divergence times (Badrane and May, 1999; Stankis et al., 1992).




I know this info doesn't simplify the discussion of the mating system, but it clears up some of the inaccuracies I've seen on the board (no offense to anyone). There seems to be a gap between the knowledge between professional mycologists and amateur mycologists (by the way I am definitely an amateur with exposure to professionals) I'm just trying to bridge some of the gap. Unfortunately, things are much more complicated than even what is touched on in the linked papers. There are other phenomenon that can occur.

I mentioned in another post that there is such a thing as incomplete mating. This happens when two monokaryons that have different alleles at one mating type locus and have the same allele for the other locus. The monokaryons fuse and form a dikaryon that is unstable and soon dies. This only occurs with one of the loci, but I don't remember which one. What I mean is that incomplete matings only happens when the A locus is the same and the B locus is different but not when the A locus is different and the B locus is the same (or vice versa since I can't remember which way it is).

Another phenomenon seen is the mating of a dikaryon with a monokaryon from which several results can occur. The first is that the nuclei from the dikaryon spread through the monokaryon. The nuclei from the monokaryon is lost. Another result is that the monokaryon's nuclei replace one of the dikaryon's and a new dikaryon is formed throughout. Another result seen is that the monokaryon takes up one of the dikaryon's nuclei to form a new dikaryon while the original dikaryon continues to grow separately.

Another point to consider is that not many mushrooms have had their mating type system studied in much detail. Of those studied, half to two thirds have the classic bifactorial mating system. Others have modifications such as one locus with multiple alleles and the other with only two. There will most likely be mushrooms with some other modifications and possibly some with completely different mating systems as well. It may well be that the mushrooms discussed most often here are the oddballs. Only time will tell.
T34

Note: I came across this post from last year that links to a relevant article from 1998.

Edited by texas34 (05/06/06 11:07 PM)

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InvisibleGio
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Re: Bifactorial mating system [Re: texas34]
    #5599856 - 05/07/06 01:31 AM (18 years, 16 days ago)

"Remarkably, a single species may have many thousands of cross-compatible mating types because the mating-type genes are multiallelic."

It almost seems like basidiomycete fungi have all sorts of genders.

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Invisibletexas34
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Re: Bifactorial mating system [Re: Gio]
    #5604581 - 05/08/06 11:16 AM (18 years, 15 days ago)

Gender is not a good word to use here, but yes there are a great many compatible mating types in this system. Even with a relatively small number of alleles at each locus, you will get a large number of possibilities. For example, if each locus has 10 alleles, then there are 100 different mating types. The linked papers say that a reasonable average is at around 100 alleles per locus, which equates to 10,000 mating types. It's actually pretty crazy when you think about it.
T34

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Invisiblefastfred
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Re: Bifactorial mating system [Re: texas34]
    #5631190 - 05/15/06 02:22 AM (18 years, 8 days ago)

Thanks for the awesome post T34. I'd always been meaning to learn more about mating types. The links you gave in another post were great.

I think that by doing some mating experiments it would be fairly easy to get a ballpark idea of the number of different alleles amongst the different strains. That should give a pretty good idea of relatedness I would think. Probably almost as useful as RFLP, without the DNA extraction and running of gels. Any thoughts on that?


-FF


--------------------
It drinks the alcohol and abstains from the weed or else it gets the hose again. -Chemy

The difference between the substances doesn't matter. This is a war on consciousness, on our right to the very essence of what we are. With no control over that, we have no need to speak of freedom or a free society. -fireseed

"If we are going to have a war on marijuana, the least we can do is pull the sick and the dying off the battlefield." -Neal Levine (MPP)

I find the whole "my drug should be legal but yours should be illegal" mindset disgusting and hypocritical. It's what George Bush and company do when they drink a cocktail and debate the best way to imprison marijuana users. -Diploid

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Invisibletexas34
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Re: Bifactorial mating system [Re: fastfred]
    #5642590 - 05/17/06 04:43 PM (18 years, 6 days ago)

FF, mating experiments would be easy once you get single spore isolates to perform your crosses. The main issue I see, which is an issue with RFLP also, is that because the strains are isolates from the wild they should only contain 2 different alleles for each mating type loci. Spores from one fruiting body will only have 4 different haplotypes (given no mutations during meiosis occurred). The population as a whole, in the environment, will have tons of different alleles but not isolated strains.

So the experiments would be straightforward in theory. Dilute spores from the two strains of your choice and spread them on water agar plates, separately of course. Check them everyday for growth, with a microscope, and transfer each single spore isolate to its own nutrient media plate. Then after getting 30 or so isolates per strain, cut out wedges from an isolate of each strain and place them together on a new plate (space them apart some). Repeat for all your isolates. You could even do different combos of your isolates also. If both alleles for each loci are different between strains, then all crosses will form dikaryons. If there are some common alleles between the strains, then you would get different ratios of successful and non-successful matings. An obscuring problem here is if any of the common alleles allow for incomplete matings. Also mutations during meiosis could lead to some new alleles being created, but you may not isolate these anyway with the massive amount of spores from a fruitbody.

If you started crossing other strains as well, then you might find some strains that share alleles. It would seem to me to be most likely that most of the isolated strains would have completely different alleles, but that's just playing on probabilities. It is definitely possible that some of the alleles are shared in the isolated strains.

In the end, mating experiments would be a bit easier than extracting DNA and the like but may lead to hard to interpret data that RFLP would avoid.
T34

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OfflineFeelers
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Re: Bifactorial mating system [Re: texas34]
    #5643146 - 05/17/06 06:54 PM (18 years, 6 days ago)

I've never really understood the mating of mushrooms from an evolutionary standpoint. Spores (monokaryons) get carried off to far away places, land and start growing. But - in order for the new colony to reproduce it needs another spore to mate with. The system obviously works in reality, but it's rather odd in theory.

Out in the wild are there loads of monokaryotic colonies eagerly awaiting spores to mate with? Do these colonies get big or are they doomed for failure? If it works it works, but I still would have thought there was a better way of going about it.

Sorry tx (this is kinda related :laugh:)

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Invisiblefastfred
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Re: Bifactorial mating system [Re: Feelers]
    #5643441 - 05/17/06 07:58 PM (18 years, 6 days ago)

The advantage of a sexual reproductive system is that you gain genetic variability which is essential for the long term survival of a species. The disadvantage is that it takes two organisms to produce offspring. If you were trapped on a desert island it would suck that you couldn't cut off a finger and grow up some offspring for company and to take care of you when you got older. Sexual reproduction usually works to the detriment of the individual, but the advantage of the species as a whole. Some spores will never fruit (reproduce) because they can't find a mate. The same is true of most species. However if there is one spore, then there are likely others that aren't too far away. Spores also clump to some extent so often 2 or more will already be stuck together when they land. Kind of like a prearranged marriage.

I'm not sure how large monokaryons get before senescence sets in or if it's longer or shorter than dikaryons. I'm sure it varies amongst species. I would think that a monokaryon would hang out until a spore happened to land on or near it. Then it would mate immediately and fruit quickly since it's growing and substrate digestion would already be mostly done. Perhaps it's an advantage to not have every square inch of the globe colonized with spores... If an organism overpopulates and consumes all it's available food it's not very good for species survival. Who knows? There are tons of factors and evolutionary genetics is one of the most hotly debated areas in science today. There are many many theories to explain the multitude of traits seen in all the species seen around the world. The problem is that there are no easy answers. The simple answer is that sexual reproduction provides valuable variation that ensures species survival, which far outweighs any advantage to the individual which would be gained by asexual reproduction.


-FF


--------------------
It drinks the alcohol and abstains from the weed or else it gets the hose again. -Chemy

The difference between the substances doesn't matter. This is a war on consciousness, on our right to the very essence of what we are. With no control over that, we have no need to speak of freedom or a free society. -fireseed

"If we are going to have a war on marijuana, the least we can do is pull the sick and the dying off the battlefield." -Neal Levine (MPP)

I find the whole "my drug should be legal but yours should be illegal" mindset disgusting and hypocritical. It's what George Bush and company do when they drink a cocktail and debate the best way to imprison marijuana users. -Diploid

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OfflineRogerRabbitM
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Re: Bifactorial mating system [Re: fastfred]
    #5643542 - 05/17/06 08:26 PM (18 years, 6 days ago)

Quote:

fastfred said:
Sexual reproduction usually works to the detriment of the individual
-FF




Dude, you've got to pick one out before closing time.
RR

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Invisiblefastfred
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Re: Bifactorial mating system [Re: RogerRabbit]
    #5643767 - 05/17/06 09:19 PM (18 years, 6 days ago)

:rofl2:


--------------------
It drinks the alcohol and abstains from the weed or else it gets the hose again. -Chemy

The difference between the substances doesn't matter. This is a war on consciousness, on our right to the very essence of what we are. With no control over that, we have no need to speak of freedom or a free society. -fireseed

"If we are going to have a war on marijuana, the least we can do is pull the sick and the dying off the battlefield." -Neal Levine (MPP)

I find the whole "my drug should be legal but yours should be illegal" mindset disgusting and hypocritical. It's what George Bush and company do when they drink a cocktail and debate the best way to imprison marijuana users. -Diploid

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OfflineFeelers
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Re: Bifactorial mating system [Re: fastfred]
    #5644489 - 05/18/06 12:08 AM (18 years, 5 days ago)

I didnt realise that spores tended to clump - that makes a bit more sense. :laugh:

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