How did torsion evolve in snails?

[Darwin123]

How did gastropods evolve torsion?
I have looked all over. There doesn’t seem to be any clear idea of what torsion does now, let alone what purpose it served long ago.
There was a hypothesis (Garstang’s hypothesis) that the veligar larvae developed torsion so they could pull their little heads into their little shells. However, that appears to have been disproved in 1985. This link tells why.
http://www.mbari.org/staff/peti/Pubs/Gastropod Torsion-A Test of Garstang's Hypothesis.pdf
“in only one case was rate of predation reduced in pretorted larvae. It therefore appears that torsion does not function defensively,…”

The inheritance of torsion involves a weird delay in phenotypic expression. The gene expresses itself in the offspring of the mother. Aside from the interesting questions in development it brings up, this suggests some very twisted natural selection.
http://science.naturalis.nl/media/280795/schilthuizendavisonnawi.pdf
“In the few species that have been characterized, chirality is determined by a single genetic locus with delayed inheritance, which means that the genotype is expressed in the mother’s offspring…Nevertheless, chiral reversal could still be a contributing factor to speciation (or to
divergence after speciation) when reproductive character displacement is involved.”

Here it is 2009, and students are still writing theses on what they don’t know about torsion. Apparently the torsion makes some snails more sensitive to ocean acidification. However, there is no evidence that it makes the snail less sensitive to anything else. So why evolve it?
http://www.escholarship.org/uc/item/39q8w7gh


It almost appears that mainstream science has given up on explaining the evolution of torsion in gastropods. Anyone have any other ideas?

 

[Borek]

This requires speculation and is on the verge of the independent research, so technically is against forum rules. That being said...

I can think of two alternatives to torsion - one is a straight cone (like orthocone), the other is a random tube (like in worm snails). The latter is present only in immobile snails, so here comes the first idea - torsion allows snails to grow the shell and stay mobile. It is much easier to carry your house in the form of a round backpack, than in the form of some shapeless, clumsy bundle.

But conical shell would be similar - it would allow growth and it would not impede mobility (too much). That's the way Turritellidae look, apparently long conical shape is not a problem (yes, the shell is twisted internally, but I am thinking just in terms of the overall shape). However - and here comes the second thought - in a twisted shell it is much easier to get out of the reach of the enemies, just retracting the body behind the first twist. Even if you meet a crab with long pincers it won't be able to get to you (unless the pincers would look like the corkscrew).

Finally, twisted shell requires less material and can be lighter, providing same defense for less effort.

 

[Ken Natton]

I happen to know someone who has done research in this very area. I don’t claim to know him well, I never actually met him, but I did have some contact with him, and I remember him talking about this very issue. My first thought, and this does not come from my contact this is my own thought, is that it is a false assumption that there has to be an evolutionary reason for it. Not all traits have a definite evolutionary purpose, some are just the result of genetic drift. It could be that snails coil because the mutation that brought it about occurred and was never selected against, until, by nothing more than chance, it had become so widespread in the species that it became fixed.

What I can tell you that did come from my contact does not answer your question and thus may be unsatisfactory to you, but I found it to be a fascinating little gem, so I’ll offer it here. The gene that controls the direction of the coil is known. Most snails, these days, are right coilers. You may see an occasional left coiler with a different allele of the gene. At one time, there was a reasonably even split of the two alleles and thus of left and right coilers. The reason that right coilers came to predominate almost certainly is a matter of nothing more than genetic drift, but having become predominant, there is now a selective advantage to being a right coiler, because it is difficult for opposite coilers to mate, and thus right coilers are far more likely to find a mate. Hence, whenever the left coiling mutation comes up, it tends to be selected against. Anyway, here’s the fascinating gem. Human beings have the exact same gene and it controls our left – right asymmetry too. Not you understand left handedness or right handedness, but the asymmetrical layout of our internal organs.

 

[zoobyshoe]

I see from the wikipedia that:

A snail's shell forms a logarithmic spiral.

http://en.wikipedia.org/wiki/Land_snail

Following the hyperlink to logarithmic spiral I find:

Spira mirabilis, Latin for "miraculous spiral", is another name for the logarithmic spiral. Although this curve had already been named by other mathematicians, the specific name ("miraculous" or "marvelous" spiral) was given to this curve by Jacob Bernoulli, because he was fascinated by one of its unique mathematical properties: the size of the spiral increases but its shape is unaltered with each successive curve, a property known as self-similarity. Possibly as a result of this unique property, the spira mirabilis has evolved in nature, appearing in certain growing forms such as nautilus shells and sunflower heads.

And:

Logarithmic spirals in nature

In several natural phenomena one may find curves that are close to being logarithmic spirals. Here follows some examples and reasons:

The approach of a hawk to its prey. Their sharpest view is at an angle to their direction of flight; this angle is the same as the spiral's pitch.[5]

The approach of an insect to a light source. They are used to having the light source at a constant angle to their flight path. Usually the sun (or moon for nocturnal species) is the only light source and flying that way will result in a practically straight line.[6]

The arms of spiral galaxies.[7] Our own galaxy, the Milky Way, has several spiral arms, each of which is roughly a logarithmic spiral with pitch of about 12 degrees.[8]

The nerves of the cornea (this is, corneal nerves of the subepithelial layer terminate near superficial epithelial layer of the cornea in a logarithmic spiral pattern).[9]

The bands of tropical cyclones, such as hurricanes.[10]

Many biological structures including the shells of mollusks.[11] In these cases, the reason may be construction from expanding similar shapes, as shown for polygonal figures in the accompanying graphic.

Logarithmic spiral beaches can form as the result of wave refraction and diffraction by the coast. Half Moon Bay, California is an example of such a type of beach.

http://en.wikipedia.org/wiki/Logarithmic_spiral

This leads me to think there is no need for a gene, or genetic explanation, for this, that it could be a purely mechanical result of the way they add material to their shells. Does a galaxy need a 'gene' to form itself into logarithmic spirals?

 

[Darwin123]

I happen to know someone who has done research in this very area. I don’t claim to know him well, I never actually met him, but I did have some contact with him, and I remember him talking about this very issue. My first thought, and this does not come from my contact this is my own thought, is that it is a false assumption that there has to be an evolutionary reason for it. Not all traits have a definite evolutionary purpose, some are just the result of genetic drift. It could be that snails coil because the mutation that brought it about occurred and was never selected against, until, by nothing more than chance, it had become so widespread in the species that it became fixed.

What I can tell you that did come from my contact does not answer your question and thus may be unsatisfactory to you, but I found it to be a fascinating little gem, so I’ll offer it here. The gene that controls the direction of the coil is known. Most snails, these days, are right coilers. You may see an occasional left coiler with a different allele of the gene. At one time, there was a reasonably even split of the two alleles and thus of left and right coilers. The reason that right coilers came to predominate almost certainly is a matter of nothing more than genetic drift, but having become predominant, there is now a selective advantage to being a right coiler, because it is difficult for opposite coilers to mate, and thus right coilers are far more likely to find a mate. Hence, whenever the left coiling mutation comes up, it tends to be selected against. Anyway, here’s the fascinating gem. Human beings have the exact same gene and it controls our left – right asymmetry too. Not you understand left handedness or right handedness, but the asymmetrical layout of our internal organs.

Genetic drift doesn't do much when the inherited feature has a large effect on the fitness of the organism. When there is "genetic drift", part of the genome in a population of organisms differentiates because of random variation with no selection. Genetic drift may explain how a feature starts developing (e.g., the "half an eye"), but is doesn't explain how a specialized feature finally develops.
Your analogy with humans doesn't work for two reasons. First, bilateral asymmetry in human beings has a big effect on the chances of survival. Second, the bilateral asymmetry in snails has a large cost to it.
The bilateral asymmetry in the mammalian body has a very strong fitness value. For example, the aorta is attached to the left ventricle and the pulmonary artery to the right ventricle. For animals with lungs, a symmetric heart would not work as efficiently.
A person with a bilaterally symmetric heart would die fairly quickly. The human heart can not function symmetrically. One side of the heart has to supply blood to the lungs, and the other side has to supply blood to the rest of the body.
The asymmetry of the brain is common to all vertebrates, but is especially enhanced in the case of the human species. The asymmetry in lower vertebrates is minor, but it seems to speed up the choices made by the vertebrate. In human beings, the two hemispheres of the brain perform different functions. Each does something the other can't do. Again, this seems to speed up certain choices. Animals that throw things have to be able to distinguish left from right very rapidly. Humans apparently evolved to throw things.
The initial steps in the evolution of asymmetry in vertebrates may have started with genetic drift. As you pointed out, the dominant chirality in the extant vertebrate body was probably "selected" in those early years. However, the the asymmetry in the extant vertebrate body is very prominent because of natural selection.
The snail pays a large price in survival because of the torsion. As shown in the links that I posted, the metabolism of the snail is stressed at the stage in development just before the torsion starts. It requires energy to twist its body. It becomes more sensitive to acidity in the water just before the torsion. So one would think there must be a compensating advantage in order for natural selection to produce such a strongly asymmetric animal.
Some freshwater snails have re-evolved the symmetric shape. In fact, these snails experience two torsions in their lives. They twist one way early in their development and then they twist the opposite way later in development. Hence, the adult has a secondary bilateral symmetry.
This second torsioning does not appear consistent with the genetic drift model. First, the second torsion only occurs in freshwater species. It makes sense that a freshwater snail would face different challenges then a marine snail. However, it makes no sense that "genetic drift" would favor marine over freshwater environments. Second, if genetic drift explained the secondary bilateral symmetry then one would expect to see snails with no torsioning at all. There should be some snails that start life bilaterally symmetric and remain that way all their lives. Instead, the snails have to go through that undo the first torsioning.
Another thing is that there are genes common to all snails committed to asymmetry in the snail. How come there are no snails that just plain don't torsion?
I haven't found any paleontologist analyze torsioning. I think paleontology could probably solve this one. When does bilateral asymmetry become common among mollusks? There are animals called rostroconchia that supposedly are "intermediate" between gastropods and bivalves. The common ancestor of rostroconchia (monoplacorans?)was probably symmetric. Are there any rostroconchs that are asymmetric?
I conjecture that whatever purpose the asymmetry served in early mollusks was the reason that the asymmetry evolved. However, I haven't found an easily understandable article discussing asymmetry in fossils.

 

[256bits]

I conjecture that whatever purpose the asymmetry served in early mollusks was the reason that the asymmetry evolved. However, I haven't found an easily understandable article discussing asymmetry in fossils.

Perhaps it would help if you would back up and make a conjecture on why a shell of any kind was a necessary adaptation. Was an the shell an adaptation to ward off predators who would not care for a crunchy snack, or as camouflage - if you are cannot be distinguished from a backgrounf of pebbles you can't be eaten., or for some other reason.

To build a shell the animal has to expend energy, which in turn would mean that more time would have to be spent searching and acquiring food. If food is plentiful then any type of shape for a shell would be adequate as long as it fulfils the requirements for having a shell. Even then, as long as predation is not extensive enough to terminate continuation of the species who cares what type of shell you have.

Any stress on the population with regards to a limited food supply, or increased predation would naturally favour those offspring who expend less energy on shell buidling.

Borek alluded to what I would think is a major reason for the spiral:

Finally, twisted shell requires less material and can be lighter, providing same defense for less effort.

and also zoobeshoe:

This leads me to think there is no need for a gene, or genetic explanation, for this, that it could be a purely mechanical result of the way they add material to their shells

although mechanical in nature there would have to be a driving force, possibly a chemical gradient from inner to outer parts of the spiral, that causes the animal to continue the spiral. I doubt though if a responsible gene(s) can be ruled out.

Just my 2 cents. I am sure you have already explored these avenues.

 

[zoobyshoe]

and also zoobeshoe:

although mechanical in nature there would have to be a driving force, possibly a chemical gradient from inner to outer parts of the spiral, that causes the animal to continue the spiral. I doubt though if a responsible gene(s) can be ruled out.

Lets say you have a band of calcium excreting glands around the part of the snail that's just inside the shell. If the snail were strong enough to support it, each new band of calcium carbonate it secretes would add to a cone sticking up in the air. But snails aren't that strong and the cone is always too heavy so it flops over. The band of glands is therefore compressed on one side and stretched on the other. The result is that each new band of calcium it secretes is wide on the upper most side and narrow on the side nearest the body. For purely mechanical, non-genetic reasons, the shell therefore comes out as a spiral when it wants to be a right regular cone. That sort of thing.

 

[Borek]

Lets say you have a band of calcium excreting glands around the part of the snail that's just inside the shell. If the snail were strong enough to support it, each new band of calcium carbonate it secretes would add to a cone sticking up in the air. But snails aren't that strong and the cone is always too heavy so it flops over.

I am not convinced - I have already mentioned Turritellidae. Their shells are even heavier than a simple cone would be (because of the additional internal structure), yet they don't flop over and some grow quite large and long. See for example http://www.gastropods.com/7/Shell_1527.shtml. Similar examples in other orders - Triseriate Auger - http://www.gastropods.com/8/Shell_1568.shtml.

The band of glands is therefore compressed on one side and stretched on the other. The result is that each new band of calcium it secretes is wide on the upper most side and narrow on the side nearest the body. For purely mechanical, non-genetic reasons, the shell therefore comes out as a spiral when it wants to be a right regular cone. That sort of thing.

While it suggests that next part is attached to the surface of the older part of the shell, it doesn't guarantee the shell to be always twisted in the same direction and symmetrically spiral - it can be as well just a clump of randomly directed twists (sorry, no idea how to better describe it in English). Something like worm snail shell growing always attached to the earlier part of the shell.

 

[Darwin123]

I am not convinced - I have already mentioned Turritellidae. Their shells are even heavier than a simple cone would be (because of the additional internal structure), yet they don't flop over and some grow quite large and long. See for example http://www.gastropods.com/7/Shell_1527.shtml. Similar examples in other orders - Triseriate Auger - http://www.gastropods.com/8/Shell_1568.shtml.



While it suggests that next part is attached to the surface of the older part of the shell, it doesn't guarantee the shell to be always twisted in the same direction and symmetrically spiral - it can be as well just a clump of randomly directed twists (sorry, no idea how to better describe it in English). Something like worm snail shell growing always attached to the earlier part of the shell.

I was originally talking about torsion in mollusks. However, check out this example of torsion in a mammal!

http://www.huffingtonpost.com/2012/10/07/sheep-with-upside-down-head-video_n_1946331.html
“Footage of Terry, a sheep that looks to have an upside-down head, has gone viral on the Internet, sparking a debate over its authenticity.
Allan McNamara, a computer technician, claims to have encountered the animal in a pasture in the north of England. He shot video of the grazing sheep, which he postulated was born with a twisted spine.
"He lives happily and has been checked by a vet to ensure he is in no pain. He can eat, sleep and do everything other sheep can," McNamara told The Daily Mail.”

Although of course I can’t be sure, I believe the story. It doesn’t really violate any law of biology. If you look at the close up shot, the bilateral asymmetry is apparent. This sheep merely has a twisted neck. It probably doesn’t have any organs out of place.

That brings up an interesting possibility. May the most recent common ancestor of all gastropods flipped upside down! For instance, a rostroconch that eats worms on the bottom of the ocean may have migrated to shallow tidal pools. Then, itmay have started floating on the surface of the water. Then, it would have been more convenient to graze with the head twisted upside down. Or the other way around.

Some freshwater snails are achiral as adults. However, they have a second torsion to counteract the first torsion. If the first torsion is a mystery, then the second torsion is even more so. If the first torsion was advantageous to the snail, or even neutral, why would natural selection undo the results of the first torsion in such an inconvenient way?

I haven’t found any studies that say a chiral shell is more mechanically stable than an achiral shell. Structural stability may explain how a coiled shell is more stable than a straight shell, hence the logarithmic curve. However, some cephalopods have lived pretty well with bilateral symmetric but achiral shells.

Maybe I should ask Terry the Sheep what the advantages are from his perspective!

 

[zoobyshoe]

I am not convinced - I have already mentioned Turritellidae. Their shells are even heavier than a simple cone would be (because of the additional internal structure), yet they don't flop over and some grow quite large and long. See for example http://www.gastropods.com/7/Shell_1527.shtml. Similar examples in other orders - Triseriate Auger - http://www.gastropods.com/8/Shell_1568.shtml.

These look flopped over to me! The alternative, which doesn't happen, is that the shell would be sticking up in the air, and each new band of material added would be the same width all way around: an untwisted, right regular cone.

While it suggests that next part is attached to the surface of the older part of the shell, it doesn't guarantee the shell to be always twisted in the same direction and symmetrically spiral - it can be as well just a clump of randomly directed twists (sorry, no idea how to better describe it in English). Something like worm snail shell growing always attached to the earlier part of the shell.

I think I understand what you're saying.

The fact the shell always hangs to the same side in most members of a species is almost surely genetic, but that's a different issue from why it spirals as it grows. It would spiral for mechanical reasons alone in my scenario even if it were flopped straight back or straight forward. You're right: if the shell went back and forth randomly, first hanging to one side, then the other, the twists would be randomly directed, but that just proves the twisting is a separate phenomenon from the preference of a side to hang toward. The direction of the twist doesn't cause the twisting. The mutants they find that twist the "wrong" way twist just as well as the normal ones.

My idea can be checked: the number and distribution of the glands that secrete the calcium could be counted and mapped, particularly in newborn and embryonic snails (before any potential atrophy from non-use develops on the "pinched" side in older snails). If the snails are born with a uniform band of these glands all around, the spiraling is mechanical. If they develop from scratch with a non-uniform band that is narrower toward the back end and one side of the snail, then it is genetic.

 

[Ken Natton]

Your analogy with humans doesn't work for two reasons. First, bilateral asymmetry in human beings has a big effect on the chances of survival. Second, the bilateral asymmetry in snails has a large cost to it...

Please believe me Darwin123, I do not suppose to argue with you, I am not in any doubt about the superiority of your knowledge. Of course, there is every chance that I explained what was explained to me very badly. Unfortunately, I no longer have access to the text of the exact conversation I had with my contact. However, I am sure that he was a bona fide researcher at a British university, and knew what he was talking about. The following links demonstrate, I hope, the accuracy of the basis, at least, of what I said, and in point of fact, the first one might actually be a better contribution to answering your original question.

http://novelty.wikispaces.com/Asymmetry+in+snail+shells+

http://en.wikipedia.org/wiki/PITX2

http://en.wikipedia.org/wiki/NODAL

 

[Darwin123]

Please believe me Darwin123, I do not suppose to argue with you, I am not in any doubt about the superiority of your knowledge./QUOTE]
Sometimes what looks like superiority of knowledge is really mediocrity of knowledge!
I don't know what is going on or I wouldn't ask! I am just an arm chair biologist. I wasn't fighting either.
I was giving my knee jerk response to the phrase "genetic drift". I am not clear what it means.

Of course, there is every chance that I explained what was explained to me very badly. Unfortunately, I no longer have access to the text of the exact conversation I had with my contact. However, I am sure that he was a bona fide researcher at a British university, and knew what he was talking about. The following links demonstrate, I hope, the accuracy of the basis, at least, of what I said, and in point of fact, the first one might actually be a better contribution to answering your original question.

http://novelty.wikispaces.com/Asymmetry+in+snail+shells+

http://en.wikipedia.org/wiki/PITX2

http://en.wikipedia.org/wiki/NODAL

Thank you for the links.

 

[Ken Natton]

I was giving my knee jerk response to the phrase "genetic drift". I am not clear what it means.

Well my understanding of the term is that it refers to changes that are the result of the same cumulative mutations that drives all change, but on which no selective pressures, either positive or negative, have acted. Like flotsam in the ocean.

 

[256bits]

The item about the sheep sparked a thought about curly hair in mammals.
and one site that popped up is this:
http://ebm.rsmjournals.com/content/223/1/1.full
in regardds to researchin molecular genetics.

Under the heading of Homeobox Genes and Curly Hair.

These findings show that genes whose ancestors are involved in making terminal appendages in fruit flies are actually making terminal skin appendages in mammals. Moreover, this is the first time that homeobox-containing genes have actually been implied in ectodermal-mesenchymal interactions in systems other than the developing limb.

and heading Wavy Hair and Fibroblast Growth Factors.

Expression of Fgf5 is detected in wild-type hair follicles and is localized to the outer root sheath during the late anagen phase of the hair growth cycle. It seems that Fgf5 in one way or another regulates the length of the anagen phase. Fgf-receptors 1–3 are involved in the regulation of skeletal growth in vertebrates. In particular, null mutant alleles of Fgfr3 cause skeletal overgrowth in mice

they do say that some factors are unknown.

what I found interesting is that a common gene between a present day fruit fly and a mammal is active and producing certain traits in each. And how a certain gene can promote functioning of what would be considered unrelalated organs such as the tooth, hair, and the skeleton.

Certain horns in mammals have a twisted nature, if not just slightly curved versus the straight or branching aspect of other horns.

Could not all the twists and torsions be traised back to a common ancestor of mammals, insects, mollusks by a common gene.

 

[zoobyshoe]

How did gastropods evolve torsion?
I have looked all over. There doesn’t seem to be any clear idea of what torsion does now, let alone what purpose it served long ago.
There was a hypothesis (Garstang’s hypothesis) that the veligar larvae developed torsion so they could pull their little heads into their little shells. However, that appears to have been disproved in 1985. This link tells why.
http://www.mbari.org/staff/peti/Pubs/Gastropod Torsion-A Test of Garstang's Hypothesis.pdf

Re-reading the first couple paragraph of this link, I see I completely misunderstood the particular torsion the term refers to. I thought it referred to the coiling of the shell alone. Instead it refers to the fact that the shells start off growing forward in the larvae, then, for some reason, become physically twisted 180° and thereafter coil backward. This means that the viscera inside are also twisted, having first formed in a 180° different orientation.

This is certainly bizarre and begs for an explanation. I apologize for going off on a tangent about logarithmic spirals, which don't really have anything to do with the torsion you want to discuss.

That said, here's a thought:

The ubiquitous land snails we have here in San Diego are extremely moisture dependent and only come out when it rains. Their shell are pretty delicate and don't seem to have any protective function against large predators (not that I know what might try to eat them). The function of their shells seems to be to retain moisture. When it's not raining they seal themselves up in them in the shadiest spot available. (It can go weeks and months here without raining.)

The ability to seal themselves up might also be important for marine gastropods as a way of retaining moisture in tidal pools at low tide, and to protect themselves from any unpleasant chemistry that develops in the water in those pools. At any rate, if land snails couldn't retract into their shells and seal them up, they would all die out fast here in this desert city. If snails can't retract and seal without the 180° torsion, then it can be seen to be vital to land snails, at least.

 

[Darwin123]

The ability to seal themselves up might also be important for marine gastropods as a way of retaining moisture in tidal pools at low tide, and to protect themselves from any unpleasant chemistry that develops in the water in those pools. At any rate, if land snails couldn't retract into their shells and seal them up, they would all die out fast here in this desert city. If snails can't retract and seal without the 180° torsion, then it can be seen to be vital to land snails, at least.

That is just it. I don't fully understand how snails retract. I know they plug up the holes with the operculum on their "tails". To do this, they have to retract totally. However, could imagine how retraction could be done using bilateral symmetric muscle sets.
The symmetric snails that I have seen seem to be able to retract fully within their shells. My father and I kept a sort of pond aquarium in our house when I was a child. One time we had a snail, which I remember as symmetric. I would usually see it retracted, hidden in its shell. I saw what I later learned was the operculum, so I think that it was fully retracted. This is all a faint childhood memory, so you should look at this as an anecdote rather than real knowledge. In any case, it was only one species of snail. I don't know how well other symmetric snails retract.
I haven't looked into the details of gastropod retraction. So I have to ask.
1) How do you know they can't retract without the 180° torsion?
2) Why can't they retract without the 180° torsion?

There is an entire order of snails, the pulmonata, that are bilaterally symmetric as adults. They torsion twice as veligar in order to come out as bilateral symmetric adults. However, I developed a reasonable (?) conjecture.
The pulmonata gastropods have a "lung" which is really a vascularized mantle. The "lung" enables them to breath air. They are mostly land snails. I conjecture that a chiral snail would have less serface area on the inner surface of their "lung." Hence, there would be a strong selection for uncoiling once the mantle got vascularized. The second torsion and the vascularization of the mantle may have evolved slowly but simultaneously.

 

[zoobyshoe]

I haven't looked into the details of gastropod retraction. So I have to ask.
1) How do you know they can't retract without the 180° torsion?
2) Why can't they retract without the 180° torsion?

I certainly don't understand the musculature, I just know what your link says (first paragraph under the heading "Introduction"), which is that, without torsion the tail is retracted first, and the head second, which leaves the head exposed at the unsealed opening. The operculum is on the tail, so sealing the shell means the tail must be the last thing retracted, not the first. Torsion is not required to retract, it's required to retract and seal.

So, by what's asserted in that paper, the snail you remember would have to be unusual to be able to retract the head first and tail second without torsion.

It mentions that something called "opisthobranch" veligers (I don't know what that means) can fully retract without torsion, but doesn't specifically say they can also seal the shell, so that's a loose end in my mind.

There is an entire order of snails, the pulmonata, that are bilaterally symmetric as adults. They torsion twice as veligar in order to come out as bilateral symmetric adults. However, I developed a reasonable (?) conjecture.
The pulmonata gastropods have a "lung" which is really a vascularized mantle. The "lung" enables them to breath air. They are mostly land snails. I conjecture that a chiral snail would have less serface area on the inner surface of their "lung." Hence, there would be a strong selection for uncoiling once the mantle got vascularized. The second torsion and the vascularization of the mantle may have evolved slowly but simultaneously.

Your conjecture sounds reasonable to me. I would want to know if they can seal their shells after the second torsion, though. If yes, how? If no, why don't they dry out when it's not raining? A land snail that can't seal its shell could survive well enough if it can get to water, or at least a moist place when the weather is dry. So a species of untorted, or re-torted, snails would survive under the condition they never had the usual reasons to completely seal themselves inside. Let's say they never stray more than 4 feet from a lake or pond. OR, they seal against moisture loss in a completely different way, say by secreting slime that dries into a moisture-proof plug. One way or another land snails have to avoid dessication.

The fact so many do seal themselves up argues there's a good reason for it. It doesn't protect against tiny predators, or against much larger predators which can swallow them whole, but it certainly must protect against near equal-sized predators, whatever those might be, maybe insects, maybe other, more aggressive snails. It is also vital to the California land snail to retain moisture. In each case where they undergo torsion to form a tight seal we'd have to examine what function it serves that snail in that environment.

Understanding exactly why torsion anatomically permits reversing the order of retraction, head first, tail second, would also certainly shed some light on why it got selected. That's not clear to me at all, I just know it was asserted in the paper that it permits it. It's also asserted torsion happened all at once, that they haven't found any partially torted fossils. That makes me wonder if the operculum preceded torsion or if it's something that broke off the main shell once the torsion mutation happened, and got refined into a nicely fitting "plug". If the operculum comes into being as a by-product of torsion, then things make a great deal more sense. If not, they remain weird.

 

[zoobyshoe]

I've been doing more reading and this idea occurred to me:

Many gastropods are hermaphrodites and can self fertilize. Let's stipulate, then, that the common ancestor of all modern gastropods was a hermaphrodite and could self fertilize. A disaster might befall its food supply. Most die. The few survivors have to disperse, to relocate as far from their fellows as possible, in order just to find enough to eat. (This is the situation in which hermaphrodites resort to self-fertilization; there's no one to mate with around.) The lucky ones find enough to eat to reproduce, but only by self fertilization.

How many generations can that go on for without the bad effects of inbreeding? Bad effects, it seems, can happen in one generation:

The effects of self-fertilization and cross-fertilization on several fitness traits were examined in the freshwater hermaphrodite snail Lymnaea peregra. Laboratory strains were established from Lake Geneva populations. Comparisons of F2 snails and their offspring showed that there are no differences in hatching time, nor in the size of young snails monitored over one month. But there was a significant difference, when the distribution of the capsule weight against the number of eggs was compared, although the effects of this on fitness are probably small. There was also a significant difference for egg production and juvenile viability over one month; the selfing snails are 94 per cent less fit for these two traits than the outcrossing.

http://www.nature.com/hdy/journal/v64/n2/abs/hdy199021a.html

Even if the ancient gastropods were dispersed for only one generation, that could have been enough to cause the torsion mutation by inbreeding (selfing). When the food supply recovered, the 'damage' would have been done.

In this scenario, torsion would not have been selected because it was advantageous in any way, or even neutral. They are worse off than before, but there was no alternative. Lots of populations survive the disadvantages of a period of inbreeding:

http://en.wikipedia.org/wiki/Inbreeding

The food supply disaster is an obviously plausible scenario, but it is only meant to stand for whatever force might cause them all to self-fertilize for one or more generations. I can think of others, like the sudden emergence of a gastropod STD: those who mate with others catch it and die, while those who self fertilize are OK. You can invent your own scenario, but forced "selfing" for whatever reason would explain why the disadvantages of torsion became standard in gastropods. Something like this would also explain the apparent suddenness and completeness of the appearance of torsion in the fossil record.

 

[Darwin123]

I've been doing more reading and this idea occurred to me:

Many gastropods are hermaphrodites and can self fertilize.

I don't think that is true. Most gastropods are hermaphrodites. However, most hermaphrodites can't self fertilize.
The male and female parts of a gastropod body are separated. They don't have little extensions that enable them to pass sperm from one part of the body to the other. I think that gastropods have to a 69 to get pregnant.
I don't think that the torsioning affects the relative position of their sex organs. The sex organs are on the part of the foot that comes after the twist that gives them chirality.
If torsioning did affect their sex act, then I would propose the opposite. Maybe torsioning came about to prevent selfing. However, I don't think the relative position of the sex organs are affected by torsioning.

Let's stipulate, then, that the common ancestor of all modern gastropods was a hermaphrodite and could self fertilize. A disaster might befall its food supply. Most die. The few survivors have to disperse, to relocate as far from their fellows as possible, in order just to find enough to eat. (This is the situation in which hermaphrodites resort to self-fertilization; there's no one to mate with around.) The lucky ones find enough to eat to reproduce, but only by self fertilization.

How many generations can that go on for without the bad effects of inbreeding? Bad effects, it seems, can happen in one generation:



http://www.nature.com/hdy/journal/v64/n2/abs/hdy199021a.html

Even if the ancient gastropods were dispersed for only one generation, that could have been enough to cause the torsion mutation by inbreeding (selfing). When the food supply recovered, the 'damage' would have been done.
http://en.wikipedia.org/wiki/Inbreeding

Inbreeding doesn't cause mutations. Inbreeding forces recessive genes to express themselves. However, the gene has to be present in the population in order to be expressed at all. Hence, the damage from inbreeding is often reversible by outbreeding.
You may be talking about a type of "founders effect" intensified by "Mendelian segregation." If there were a bunch of snails that had evolved to do selfing, and only selfing, then the different lines of snails would become homozygous for one traite or the other (the segregation). Then, if a mass extinction exterminated all lines except the homozygous "torsioned line", then what would be left are snails that only self and are all torsioned.
The problem is the same. Extant gastropods don't self. There doesn't seem to be a good reason for hermaphrodites to self all the time. There is no advantage to constantly selfing without end.
As you pointed out, selfing is a good thing after a disaster. However, it is a bad thing for most of the time. Furthermore, it doesn't cause mutations.

The food supply disaster is an obviously plausible scenario, but it is only meant to stand for whatever force might cause them all to self-fertilize for one or more generations. I can think of others, like the sudden emergence of a gastropod STD: those who mate with others catch it and die, while those who self fertilize are OK. You can invent your own scenario, but forced "selfing" for whatever reason would explain why the disadvantages of torsion became standard in gastropods. Something like this would also explain the apparent suddenness and completeness of the appearance of torsion in the fossil record.

My question was whether torsioning really appeared suddenly. I wonder if it is found in basal gastropods. This is why I asked about rostroconchs. The basal gastropod may have been a rostroconch rather than a full blown snail. So maybe there is a gradual transition to torsioning among the rostroconchs.

 

[zoobyshoe]

Inbreeding doesn't cause mutations. Inbreeding forces recessive genes to express themselves. However, the gene has to be present in the population in order to be expressed at all. Hence, the damage from inbreeding is often reversible by outbreeding.

This is what I meant. "Mutation" was the wrong term, sorry.

You may be talking about a type of "founders effect" intensified by "Mendelian segregation." If there were a bunch of snails that had evolved to do selfing, and only selfing, then the different lines of snails would become homozygous for one traite or the other (the segregation). Then, if a mass extinction exterminated all lines except the homozygous "torsioned line", then what would be left are snails that only self and are all torsioned.

"Founder's Effect" sounds right. However, I was trying to sketch a situation where the "founders" were forced to self long enough to bring the torsion genes out, but which did not wipe out their ability to have sex with others. They wouldn't be stuck selfing forever. But once they could get back with their own kind, all of them were torted, because this was the first latent tendency waiting to come out in their selfing, just as lower egg production and poor juvenile viability seems to be the first tendency to come out from selfing in the species of snails in the study I linked to. There would have been no normal, untorted ones left to outbreed with because all surviving gastropods would have been forced to self enough times to bring the torsion gene out.

The snails that were forced to self in the study I linked to all developed the same 2 problems: "There was also a significant difference for egg production and juvenile viability over one month; the selfing snails are 94 per cent less fit for these two traits than the outcrossing." I am assuming if we put the offspring of all these snails who had been forced to "self" together, these offspring could resume breeding as normal, fertilizing each other, but that the two bad traits: lower egg production and much poorer juvenile viability, would be expressed in all the successive generations.

The modern pulmonate can self fertilize:

The majority of pulmonates in fresh water are hermaphrodites and are capable of self-fertilization as well as cross-fertilization with other individuals. As a result, any pulmonate entering a new body of water can establish a considerable population of that species in a short time.

http://www.britannica.com/EBchecked/topic/226777/gastropod/35712/Ecology-and-habitats

They do this when they happen to end up where there's no one to mate with. But, as the study demonstrates, even one generation of this can bring bad genes to the fore. I'm just asking you to consider a "founder" gastropod which could self: hundreds of millions of years ago; the original gastropod, the only gastropod game in town. (Gastropods which can't self today would have evolved that later, and yet still be descendants of the "founder" population.) Selfing is an advantage because individuals that happen to become isolated can still produce offspring, and the expression of bad traits will most likely be corrected later when there's a rejoining to the original population. The gastropod disaster, however, throws a monkey wrench into that system by forcing all the individuals to self for some brief period, and the result is the offspring all express the "bad" genes for torsion.

Since all gastropods tort, even slugs (who have no shell anymore!) it makes more sense in my mind to think they all got it from a "founder" population which could self and was forced to self, and then couldn't correct the resultant deformity with outbreeding, because there were no non-deformed, non-torted members of its kind left to breed with. It's a disadvantageous deformity. In the millions of years since it first appeared all the gastropods have evolved a myriad of excellent accommodations to it, but they don't seem to be able to get rid of it, just as the Hapsburgs would never get rid of their chin if they kept breeding with other Hapsburgs (or outbreeding with Lenos, for that matter).

The alternative to this way of thinking, as far as I can see, is that there were many, many different kinds of gastropods and they all, at different times, and independently of each other, all found a separate advantageous reason to tort such that now, there are none who don't tort. Seems a stretch. Some tort and then de-tort, but the detorsion is a separately evolved correcting mechanism that came later. If you're a gastropod, you tort (according to wiki). If that doesn't trace back to a common "founder" population, I would think we'd have a lot of gastropods today that just never tort at all.

(Speaking of wiki, it describes the mechanism of torsion:

There are two different developmental stages which cause torsion. The first stage is caused by the development of the asymmetrical velar/foot muscle which has one end attached to the left side of the shell and the other end has fibres attached to the left side of the foot and head. At a certain point in larval development this muscle contracts, causing an anticlockwise rotation of the visceral mass and mantle of roughly 90˚. This process is very rapid, taking from a few minutes to a few hours. After this transformation the second stage of torsion development is achieved by differential tissue growth of the left hand side of the organism compared to the right hand side. This second stage is much slower and rotates the visceral mass and mantle a further 90˚. Detorsion is brought about by reversal of the above phases.

http://en.wikipedia.org/wiki/Torsion_(gastropod))

So, the gene, or genes, for torsion seem to simply instruct the veliger to unilaterally tense up one muscle at a certain time. The article implies, as I read it, that the longer, second stage of asymmetrical tissue growth happens naturally as a result of this constriction (meaning, there's no separate genetic instructions for the differential tissue growth). Do you read it this way?

My question was whether torsioning really appeared suddenly. I wonder if it is found in basal gastropods. This is why I asked about rostroconchs. The basal gastropod may have been a rostroconch rather than a full blown snail. So maybe there is a gradual transition to torsioning among the rostroconchs.

One thing that's clear to me from a couple days of reading is that the subject of snails and slugs is an infinity I never suspected. I haven't looked into the subject of the fossil record and I'm content to take the assertion about the suddenness of the appearance of torsion in that paper as fact, mostly just to limit what I have to think about. If you dig up info that really explodes that assertion, then I'll have to think about it.

Regardless, I just googled and found this book:

http://books.google.com/books?id=nm...ed=0CFQQ6AEwCQ#v=onepage&q=tergomyans&f=false

Which says: "We cannot directly observe torsion on fossils, so it is difficult to demonstrate that any fossil is a gastropod…"

In any event, your original question is a damned good one and I'm really just trying to resolve the cognitive dissonance it produced in me in my clumsy, un-rigorous way.

 

[Darwin123]

Which says: "We cannot directly observe torsion on fossils, so it is difficult to demonstrate that any fossil is a gastropod…"
In any event, your original question is a damned good one and I'm really just trying to resolve the cognitive dissonance it produced in me in my clumsy, un-rigorous way.

Thank you.
You have cognitive dissonance, too? I myself try not to think about it!
I never thought that one could directly see the torsion in mollusks. However, we can surely see chirality in the shells of mollusks.
Torsion starts with the tightening of a muscle. No fossil there. Torsion would produce chiral shells.
Cephalopods in general have achiral shells while marine gastropods have chiral shells. This rule is true 100% of the time. However, it is a good guide. The reason that few cephalopods are chiral is that they don't torsion.
Cephalopod shells often spiral, but are achiral. Amateur paleontologists (like myself) usually distinguish cepholopod fossils from gastropod fossils by their chirality. It is easy to tell, even for an amateur, which shells have chirality and which don't.
That is what I was hoping some paleontologist would write about chirality. I heard a lecture about mollusks which claimed that there was gradual evolution going from a rostroconch to bivalves and gastropods. Since most bivalves have achiral shells, while most gastropods have chiral shells, I conjectured that the transition from rostroconchs to gastropods showed a stepwise development of chiral shells.
I am sorry I didn't catch the fellow and ask him more questions. That was a fascinating lecture.

 

[Darwin123]

This is what I meant. "Mutation" was the wrong term, sorry.

"Founder's Effect" sounds right. However, I was trying to sketch a situation where the "founders" were forced to self long enough to bring the torsion genes out, but which did not wipe out their ability to have sex with others. They wouldn't be stuck selfing forever. But once they could get back with their own kind, all of them were torted, because this was the first latent tendency waiting to come out in their selfing, just as lower egg production and poor juvenile viability seems to be the first tendency to come out from selfing in the species of snails in the study I linked to. There would have been no normal, untorted ones left to outbreed with because all surviving gastropods would have been forced to self enough times to bring the torsion gene out.

The snails that were forced to self in the study I linked to all developed the same 2 problems: "There was also a significant difference for egg production and juvenile viability over one month; the selfing snails are 94 per cent less fit for these two traits than the outcrossing." I am assuming if we put the offspring of all these snails who had been forced to "self" together, these offspring could resume breeding as normal, fertilizing each other, but that the two bad traits: lower egg production and much poorer juvenile viability, would be expressed in all the successive generations.

The modern pulmonate can self fertilize:



http://www.britannica.com/EBchecked/topic/226777/gastropod/35712/Ecology-and-habitats

They do this when they happen to end up where there's no one to mate with. But, as the study demonstrates, even one generation of this can bring bad genes to the fore. I'm just asking you to consider a "founder" gastropod which could self: hundreds of millions of years ago; the original gastropod, the only gastropod game in town. (Gastropods which can't self today would have evolved that later, and yet still be descendants of the "founder" population.) Selfing is an advantage because individuals that happen to become isolated can still produce offspring, and the expression of bad traits will most likely be corrected later when there's a rejoining to the original population. The gastropod disaster, however, throws a monkey wrench into that system by forcing all the individuals to self for some brief period, and the result is the offspring all express the "bad" genes for torsion.

So pulmonates can self fertilize?! That is very interesting!
The reason this is interesting is that pulmonates are also the only order of snails that are achiral as adults. The adult pulmonate is bilaterally symmetrical. This is the group that I told you about before which has to tort twice to be symmetrical as adults.
This leads me to think the first tort is supposed to prevent selfing. If no other snails but the pulmonates are symmetric as adults, and if no other snails then the pulmonates are symmetrical as adults, then it suggests that the chiral asymmetry in other orders may have an advantage in preventing selfing.
Selfing is usually a disadvantage since it brings recessive genes out. So usually organisms evolve in such a way to discourage it. It also has the same disadvantage as asexual reproduction with regards to parasites. An inbred population is usually vulnerable to parasites, which reproduce and evolve fast. Outbreeding allows the population to adapt to new varieties of parasites. So both asexual reproduction and self fertilization are selected against if there are a lot of parasites around.
Let us suppose that the bilateral asymmetry prevents self fertilization.
Since cephalopods are monosexual (do I have the phrase right?), they never had a problem with selfing. Just by being one sex, they avoid self fertilization. So they never evolved a tort at all.
Maybe the first tort evolved among basal snails in an environment which was full of parasites. Self fertilization makes a population vulnerable to parasites. So snails that didn't tort self fertilized themselves, and the descendents tended to die due to parasites. Therefore, the first tort evolved.
Maybe the second tort evolved in an environment where parasites were no so common. Maybe it was hard to find other snails in that environment. Pulmonata are well adapted to dry land. Maybe dry land is more free of parasites. However, it is harder to find snails on dry land. So selfing is an advantage for two reasons.
Therefore, there are two reason to self fertilize. So the second tort evolved to permit the snails to self fertilize. They twisted themselves back in shape again to mate with themselves.
I wish I knew mollusk anatomy better to evaluate this conjecture. However, thank you for starting me on the way.

 

[Darwin123]

This is what I meant. "Mutation" was the wrong term, sorry.
The modern pulmonate can self fertilize:



http://www.britannica.com/EBchecked/topic/226777/gastropod/35712/Ecology-and-habitats

They do this when they happen to end up where there's no one to mate with. But, as the study demonstrates, even one generation of this can bring bad genes to the fore. I'm just asking you to consider a "founder" gastropod which could self: hundreds of millions of years ago; the original gastropod, the only gastropod game in town. (Gastropods which can't self today would have evolved that later, and yet still be descendants of the "founder" population.) Selfing is an advantage because individuals that happen to become isolated can still produce offspring, and the expression of bad traits will most likely be corrected later when there's a rejoining to the original population. The gastropod disaster, however, throws a monkey wrench into that system by forcing all the individuals to self for some brief period, and the result is the offspring all express the "bad" genes for torsion.

I appear to be wrong about torsion preventing self fertilization. Self fertilization is twisted.

Primitive snails release their sperm in the water. Torsioning can not prevent self fertilization in such animals because the exact position of the sex organs aren’t necessary for self fertilization.

http://www.ucmp.berkeley.edu/taxa/inverts/mollusca/gastropoda.php
“Most gastropods have separate sexes but some groups (mainly the Heterobranchia) are hermaphroditic. Most hermaphroditic forms do not normally engage in self-fertilization. Basal gastropods release their gametes into the water column where they undergo development; derived gastropods use a penis to copulate or exchange spermatophores and produce eggs surrounded by protective capsules or jelly (see Busycon spiratus photo below).”


Most snails, including pulmonates, have mechanisms to prevent self fertilization. Self fertilization is rare even amound pulmonates.

http://www.mhhe.com/biosci/pae/zoology/houseman/student/workbook/molluscawb.mhtml
“Being monoecious organisms, snails have separated the events of sperm transfer and fertilization to prevent self-fertilization. What complicates the organization of their reproductive system is that torsion has resulted in lost structures and other structures are shared by the female and male reproductive systems.”

http://www.olympusmicro.com/micd/galleries/moviegallery/pondscum/gastropoda/aquatics

“Many species are hermaphroditic, but avoid self-fertilization by transferring sperm in packets.”

I found a site which lists some current theories of how torsioning got started. I currently have an interest in the theory that it stabilizes the orientation of larva in a marine environment. Snails that live on land have less reason to torsion. Snails that live on land and have no larval stage, like the pulmonata, have no reason to torsion. Maybe that is why the pulmonata developed the second tort.

http://www.nuc.edu.ng/nucsite/File/UNN%20Inaugural%20Lectures/47th%20Lecture.pdf
“Torsion: The Principal diagnostic criterion for the members of the class gastropoda is torsion. The process of torsion was thought to be due to two different gradual adaptive processes:
(a) To regulate stabilization of the larval equilibrium
(b) To regulate balancing posture in the plantigrade stage. The process of torsion therefore regulates differential growth processes e.g. shifting the mantle cavity into the anterior position. The mantle or shell sinus already existing appears to
be a prerequisite for the survival of such torted animals in not shedding their waste products towards the inhalant currents. The regulative growth also includes the development of the right pallial organs and the right dorsoventral retractor muscle.”

 

[zoobyshoe]

The main drift of my thinking has been that it needn't have been an advantage or neutral. It could have represented a definite down grade that, never-the-less, wasn't bad enough to kill them off. The first torted gastropods may, in fact, have had a much poorer quality of life than their immediate and more healthy predecessor, but were able to survive regardless.

The second drift of my thinking is that it was advantageous but for reasons that are lost to history. We would have to know the exact physiology of the first gastropod, and the features of its environment, to understand why torsion was an advantage, things that can't be inferred from modern examples, which have branched out so much that no feature of modern gastropods can be confidently ascribed to the first ancient ones, except torsion.