Unsolved Problems in Reef Chemistry #1 Snail Shell Coiling

Randy Holmes-Farley

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I decided to start a series of threads that detail some problems in Reef Chemistry/Biology that are unsolved.

Not problems in how to do something as a hobbyist, but fundamental issues that remain unsolved by scientists, and that are clearly impacting your reef aquarium.

Today's Problem is the coiling of snail shells.

Snail shells of a single species typically coil in one direction only. Sometimes mutants are found that coil in the other direction, but in general, it is not a random event for each snail: each member of a single species all coil with the same chirality, or the "direction" of coiling. A different species may coil in the opposite direction while living in a similar location.

Here's a drawing to understand the issue:

https://en.wikipedia.org/wiki/Gastropod_shell

Neptunea_-_links%26rechts_gewonden.jpg



Why does this happen?

Scientists have studied it for hundreds of years. Ideas have come and gone.

The answer is still not understood on a molecular level, but it likely relates to the fact that at the most fundamental level, many biomolecules have a "handedness" or directionality inherent to them. That at least allows for a preferred directionality that can be genetic, rather than environmental.

Every amino acid, for example, can exist in at least two mirror image forms. The drawings below show what that means:

https://www.vanderbilt.edu/vicb/images/discoveries images 2010/reverse_polarity_fig_4.jpg

reverse_polarity_fig_4.jpg


All living creatures on earth use a particular handedness (the same handedness) of the amino acids in their proteins, not a random mix. Hence, every protein has a particular "handedness" based on its composition.

This sort of molecular mechanism for "handedness" can then potentially propagate up to the level where you can see it in your tank, as in a coiled snail shell.

A paper from a couple of years ago makes some suggestions and summarizes the history of the issue:

Snail Chirality: The Unwinding
https://www.sciencedirect.com/science/article/pii/S0960982216300604

"Unravelling the precise mechanisms of how body and shell coiling are effected is probably a long way away, given the lack of genetic tools in snails. "

I won't give any more discussion to the paper's ideas except to say they suggest it happens at the very earliest stage of development. That somehow the stage with just a few cells gets oriented in a way that causes the coiling of the shell (and the internal body layout) in a particular direction. Beyond that, it is heavy biochemistry that you can read for yourself if interested. :)

Just something to think about as you watch your aquarium. :)
 
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Interesting....never really thought about this.

The couple things that immediately came to mind:


Coriolis force (effect) - that is, detecting the rotation of the earth and because of their biology, curling accordingly. This would be relative simple to test by taking a species in the northern hemisphere and breeding it in the southern hemisphere. If true, it would curl in the opposite direction.


Dextrorotation and levorotation - Biochemical ability to detect light rotation that's tied into the genes that cause shell curling. Way to test....raise snails in total darkness and see which way they curl.


Darwin effect - I'm scratching my head on this one trying to think if there would be an advantage to direction of the curl....survival of the fittest. Do they taste different? Is one curl direction more likely to be poisonous?


Now I'll go back and read the links.
 
I doubt it has anything to do with chirality on a molecular level. Most organisms break symmetry early in their development, e.g. liver on the right in humans. I assume that it has to do with intercellular signalling that occurs during differentiation.
 
I doubt it has anything to do with chirality on a molecular level. Most organisms break symmetry early in their development, e.g. liver on the right in humans. I assume that it has to do with intercellular signalling that occurs during differentiation.

And how do you propose that ends up with a single chirality?
 
I doubt it has anything to do with chirality on a molecular level. Most organisms break symmetry early in their development, e.g. liver on the right in humans. I assume that it has to do with intercellular signalling that occurs during differentiation.

Following up more, this area too seems to be hot for study, and, I contend, may ultimately derive from molecular chirality since there may be no other inherent chirality at the gene level:

Cell Chirality Drives Left-Right Asymmetric Morphogenesis
https://www.frontiersin.org/articles/10.3389/fcell.2018.00034/full

"To explain the molecular basis of LR asymmetric development, Walport proposed the “F molecule” hypothesis (Brown and Wolpert, 1990). In this hypothesis, the F molecule is chiral and can be arranged along the anterior-posterior and dorsal-ventral axes. An object is chiral if it cannot be superposed onto its mirror image. By virtue of these properties, the F molecule can direct the LR axis based on its chirality. This idea is supported by findings on the molecular mechanisms of LR asymmetric development in mouse."

"Thus, chirality at the cellular level may broadly contribute to LR asymmetric development in various invertebrate species. Recently, cell chirality was also reported for various vertebrate cultured cells, and studies suggested that cell chirality is evolutionarily conserved, including the essential role of the actin cytoskeleton. Although the biological roles of cell chirality in vertebrates remain unknown, it may control LR asymmetric development or other morphogenetic events. The investigation of cell chirality has just begun, and this new field should provide valuable new insights in biology and medicine."

"The investigation of cell chirality is still in its infancy, and many questions, especially about the molecular mechanisms of its formation and biological functions, are still open. Answering these questions will add valuable insight for studies in biology and medicine in the near future."
 
And how do you propose that ends up with a single chirality?
It doesn't end up with a single chirality. As you say, there are occasionally mutants with the opposite chirality. Would you suggest that the biomolecules in these mutants must also have the opposite chirality? Obviously, amino acids and sugars are very unlikely to have opposite chirality in these mutants.

The complexity of the intracellular signalling that occurs during differentiation is pretty amazing. The chemical environment cells on the left may differ from cells on the right during differentiation causing differences in the tissue structures that produce the shell.
 
It doesn't end up with a single chirality. As you say, there are occasionally mutants with the opposite chirality. Would you suggest that the biomolecules in these mutants must also have the opposite chirality? Obviously, amino acids and sugars are very unlikely to have opposite chirality in these mutants.

The complexity of the intracellular signalling that occurs during differentiation is pretty amazing. The chemical environment cells on the left may differ from cells on the right during differentiation causing differences in the tissue structures that produce the shell.

I don't have all the answers, but the above article does suggest it derives from molecular chirality early on (maybe added after you posted), and I cannot see any other viable mechanism that is not environmental in some fashion.

I don't actually see any problem thinking that a single mutant might induce the opposite chirality. A single mutant can easily greatly change the conformation of a protein, and I would not make the assumption that an exact mirror image is needed to induce the opposite growth since only 2 chiral orientations of growth are possible.
 
I don't have all the answers, but the above article does suggest it derives from molecular chirality early on (maybe added after you posted), and I cannot see any other viable mechanism that is not environmental in some fashion.

I don't actually see any problem thinking that a single mutant might induce the opposite chirality. A single mutant can easily greatly change the conformation of a protein, and I would not make the assumption that an exact mirror image is needed to induce the opposite growth since only 2 chiral orientations of growth are possible.

I don't have the answers either. I just find the assertion that the shell chirality arises from the chirality of the fundamental biomolecules a bit presumptive (and dubious). There could be a breaking of symmetry at a cellular level that induces the chirality in the shell as well. Imagine a mass of undifferentiated cells. One cell spontaneously (and randomly) starts secreteing a signally chemical on one side of the cell. This could then induce a cascade of signals in adjacent cells that end up being chiral in nature.

To me, this seems more plausible than a genetic mutation that results in opposite chirality in a protein (or polysaccharide) (without altering the proteins/saccharide function) despite fixed chirality in amino acids (or sugars).
 
I don't have the answers either. I just find the assertion that the shell chirality arises from the chirality of the fundamental biomolecules a bit presumptive (and dubious). There could be a breaking of symmetry at a cellular level that induces the chirality in the shell as well. Imagine a mass of undifferentiated cells. One cell spontaneously (and randomly) starts secreteing a signally chemical on one side of the cell. This could then induce a cascade of signals in adjacent cells that end up being chiral in nature.
.

Since the answer is not known, we'll just have to disagree. :)

On the face of it though, I cannot imagine anything that isn't chiral inducing single enantiomer chirality. Where does it initially come from? I contend molecules are the only genetically controlled way. Environmentally, it could theoretically be things like coriolis, but environmental issues do not seem to fit species-wide similarity.
 
Since the answer is not known, we'll just have to disagree. :)

On the face of it though, I cannot imagine anything that isn't chiral inducing single enantiomer chirality. Where does it initially come from? I contend molecules are the only genetically controlled way. Environmentally, it could theoretically be things like coriolis, but environmental issues do not seem to fit species-wide similarity.

Look into cellular automata. Plenty of examples of these breaking symmetry despite the structure/rules being agnostic about symmetry.
 
Since the answer is not known, we'll just have to disagree. :)

On the face of it though, I cannot imagine anything that isn't chiral inducing single enantiomer chirality. Where does it initially come from? I contend molecules are the only genetically controlled way. Environmentally, it could theoretically be things like coriolis, but environmental issues do not seem to fit species-wide similarity.
To be fair, I have not read all those articles, but I have looked into this previously. Is there any evidence that there is a an opposite enantiomer of any biomolecules in snails with opposite spirals? If not, then it is also a presumption that there is any single enantiomer chirality involved.
 
Look into cellular automata. Plenty of examples of these breaking symmetry despite the structure/rules being agnostic about symmetry.

I'm not sure I understand those words :D, but I stand by my statement that chirality is not spontaneous and that molecular chirality is one of the current theories about how whole organisms develop chirality.

FWIW, this fairly current article (2016) would seem to suggest that your suggestion of chiral body plans (liver on right) not being due to molecular chirality is not rejected by the literature:

Cell chirality: its origin and roles in left–right asymmetric development
https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5104503/

"These studies suggested that molecular chirality directs whole-cell chirality. "

"Given that many types of cells from various organs and organisms show cell chirality, mechanisms driven by cell chirality might be a common platform for the development of organ-intrinsic LR asymmetry.""
 
To be fair, I have not read all those articles, but I have looked into this previously. Is there any evidence that there is a an opposite enantiomer of any biomolecules in snails with opposite spirals? If not, then it is also a presumption that there is any single enantiomer chirality involved.

I think you are making a big assumption that only a fully opposite enantiomer will induce a shift to the only other form available. Like a knife balanced on an edge, it must fall one way or the other. What pushes it one way need not be the exact opposite chemistry of what pushes it the other. :)
 
I'm not sure I understand those words :D, but I stand by my statement that chirality is not spontaneous and that molecular chirality is one of the current theories about how whole organisms develop chirality.

FWIW, this fairly current article (2016) would seem to suggest that your suggestion of chiral body plans (liver on right) not being due to molecular chirality is not rejected by the literature:

Cell chirality: its origin and roles in left–right asymmetric development
https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5104503/

"These studies suggested that molecular chirality directs whole-cell chirality. "

"Given that many types of cells from various organs and organisms show cell chirality, mechanisms driven by cell chirality might be a common platform for the development of organ-intrinsic LR asymmetry.""

I'm not saying that molecular chirality can't give rise to cellular (and greater) chirality. I'm saying that there are other mechanisms by which chirality can arise spontaneously. Further, even if the cells are chiral, that need not be the mechanism by which larger structures develop chirality.

Cellular automata are mathematical constructs governed by simple rules that give rise to all sorts of organized structures. Some of these structures may break symmetry. From a intercellular signalling perspective, I think you just need a few gradients of different chemicals along different axes to develop chirality spontaneously (don't hold me to that as I haven't gone through the geometry yet).
 
I think you are making a big assumption that only a fully opposite enantiomer will induce a shift to the only other form available. Like a knife balanced on an edge, it must fall one way or the other. What pushes it one way need not be the exact opposite chemistry of what pushes it the other. :)

I am not making that assumption. If we are talking about proteins, then basically all amino acids have the same orientation. How is an opposite chirality protein going to be consistently produced when it relies on a point mutation of an opposite chirality amino acid? There would need to be a unique codon for the opposite enantiomer. I am not aware of unique codon for specific enantiomers of amino acids.
 
The coriolis effect would be way too small of a force to dictate the direction that it would grow. The toilet bowl thing is a myth too I'm pretty sure. I would like to mention the golden rule if someone hasn't already
 
The coriolis effect. Have you checked a snail below the equator?
 

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