Embryology Biology 441 Spring 2012 Albert HarrisSymmetry
A brief but necessary digression about the concept of symmetry, and "symmetry breaking".What key ideas about symmetry do you need?#1) There are more kinds of symmetry than reflection symmetry. Displacement symmetry (somites, for example; and ions in crystals; Many examples in two and three dimensions; stripes & dots on animals.
Dilation symmetry (magnification or shrinkage) Rotation symmetry (propellers, snail shells) Combinations can occur between two or more kinds of symmetry: ...between rotational and dilation symmetry in snail shells: Magnify by 50 % equivalent to rotation by some number of degrees ...between reflection and displacement in "glide reflection" symmetry. Given a letter of the alphabet, or a drawn shape, can you state which kinds of symmetry it has, or which combinations? A, B etc N, S etc. H, I, 8 etc. * etc. #2) Weyl's way of defining symmetries: "...If there is something you can do to it, and it looks the same.. (reflect it, displace it, rotate it...distort it...anything that leaves it looking the same, then the thing has reflection symmetry, displacement symmetry, rotational symmetry) Notice this allows you to invent/discover your own new kinds of symmetry, that are useful for comprehending how embryos create shapes. Most (all?) embryological shape changes are reductions in some symmetry. Somite formation reduces from infinite displacement symmetry to displacement symmetry by just certain amounts. Likewise, consider a spherical embryo loosing infinity minus one planes of reflection symmetry. (Is it helpful to consider conjoined twins as having lost infinity minus two planes of reflection symmetry?) #3) Not just objects have particular symmetries, so do forces and processes. In the clock & wavefront theories of somite formation, a timedisplacement symmetry produces a reduced spacedisplacement symmetry (somites) If the forces causing a shape have spherical symmetry (the same strength in all directions), then what shape will these forces create? In order to make a less symmetrical shape, what property has to change? #4) Curie's Law, and embryological mechanisms that evade this law. Effects can't have less symmetry than their causes. (no problem gaining symmetry) For example, can spherically symmetrical forces produce square shapes? What shapes can they produce. If a crystal structure has inversion symmetry with respect to electrical charges, can it be piezoelectric? No, it can't. And the converse is also true! Piezoelectric properties are caused by NOT having inversion symmetry of charge. This is how Curie, a codiscoverer of piezoelectric effects, got interested in how to relate phenomena to symmetries, maybe a bigger discovery than codiscovering radium! Oh, yeah? Let's see somebody destroy a city with symmetry? ...besides Le Corbusier! #5) How to evade Curie's Law (become less symmetrical, but not randomly) is THE central problem of developmental biology. This opinion is shared by nearly all biologists who collect books on symmetry, which is hardly any.
What methods do embryos use to make themselves less symmetrical, but without becoming random?
This is a central question of Embryology;
arguably THE central question, How to develop the same asymmetries every time, such as the right/left asymmetry? ___________________________________________________________
Why are oocytes and early embryos spherical? It's because their shapes are caused by counterbalances of highly symmetrical forces. The outward fluid pressure inside an oocyte, has equal strength in all directions (= is spherically symmetrical). The contractile and elastic tension in its surface also have equal strength in all directions (parallel to this surface) Symmetrical forces create shapes with the same symmetries as the forces
Most people are not used to thinking of forces as having symmetries,
but they do. Even algebraic equations and laws of nature have symmetries. For us, the important fact is that to create a less symmetrical shape, some of the forces would have to be stronger in some particular direction. (in other words, some force would have to be less symmetrical then the shape of the embryo can also become less symmetrical, and have the same reduced symmetry as the force that causes this change.) When anything becomes less symmetrical, that is called "symmetry breaking", although it doesn't necessarily mean anything is breaking, just that something is becoming less symmetrical than it had been until then. ________________________________________________________________ Example:
Visualize an ordinary dinner plate and a drinking glass;
they both have axial symmetry. They don't have the same shape; but they do have the same symmetry. If you drop a glass or a plate on a hard surface, it probably will break into many asymmetrical pieces. That is a crude example of symmetry breaking, and not difficult. Much more difficult would be to break a glass or a plate so that it broke into two symmetrical halves; and even more difficult would be to break them consistently into slightly different right and left halves. Embryos manage to break symmetry with consistent results. ___________________________________________________________________________________ First concept: How to define symmetry?
Consider the capital letter A. The letters B, C, D and E all have horizontal planes of reflection symmetry.
? What other letters have planes of reflection symmetry (I count a total of 17)? What about the letter H?
H has a vertical plane of reflection symmetry,
and also has a horizontal plane of reflection symmetry. The letter "I" also has two perpendicular planes of reflection symmetry.
What symmetries do the numbers 8 and 0 have?
A circle has an infinite number of planes of reflection symmetry,
(Notice the contrast with H, 8 and 0, and squares,
all of which have nonequivalent planes of symmetry. Most embryos start out with an infinite number of planes of symmetry, like a cone, or a hemisphere, dinner plate or water glass. ("axial symmetry") And then special mechanisms reduce = "break" symmetry. Mammal oocytes come close to having spherical symmetry, but for most kinds of animals, the yolk is more concentrated near the vegetal pole from the very beginning, so they never have more than axial symmetry. Formation of the grey crescent breaks axial symmetry and replaces it with one plane of reflection symmetry. This amounts to the destruction of infinityminusone planes of reflection symmetry. Second big concept: There are more kinds of symmetry than just reflection. Please consider the symmetry of the letters S, N and Z.
They all have twofold rotational symmetry. The internal microtubule structures of cilia and flagella have 9fold rotational symmetry, but do not have reflection symmetry. Vertebrate embryos use this lack of reflection symmetry to control creation of RightLeft differences, (very much like the "Right Hand Rule" in the physics of electromagnetism.) Mutants (including mutant humans) who lack beating flagella nevertheless develop rightleft asymmetry more kinds of symmetry
Consider the symmetry of the following pattern @@@@@@@@@@@ A line _______________________________________ can be said to have infinite displacement symmetry, in that it it looks the same for any (small or big) amount of displacement. but this line  only has displacement symmetry for displacements of a certain distance. (a sine wave is another example). When an embryo becomes segmented, & when it forms somites, Then it is "breaking" displacement symmetry. QUESTION FOR CLASS DISCUSSION: Turing's reactiondiffusion mechanism serves to break what kind of symmetry? Even more kinds of symmetry
Another kind of symmetry is dilation symmetry.
Snail shells have combined rotation and dilation symmetry;
Certain phenomena look the same at different size scales. I recently bought an applied physics book titled "Scaling", which is about variations in the extent to which physical phenomena (explosions, for example) produce the same shapes ("mushroomshaped clouds" for example) over wide ranges in sizes. SelfSymmetry of Fractals is also amusing. (but please don't let yourself get carried away with fractals & chaos theory; neither is a hundredth as important as the principles of symmetry). You can invent new kinds of symmetry, such as reflection in a distorting mirror A Third big concept, related to symmetry Stability different kinds of stability
Divergence away from a stable state: Can break symmetry. Convergence toward a stable state (can create shape, & create symmetry) Is best achieved by negative feedback, homeostasis
You will not be tested on this physics, about conservation laws. For every symmetry, there is a conservation law. Displacement in time → behave the same→ Conservation of Energy Displacement in space → behave the same→ Conservation of Momentum Rotation of angle → behave the same → Conservation of Angular Momentum Change in quantum mechanical phase → → Conservation of Electrical Charge These facts were discovered by Prof Emmy Noether, of Bryn Mawr College, one of the 3 Quaker colleges in the suburbs of Philadelphia.
To quote Richard Feynman: What about symmetry with respect to mirror image reflection? No! Experiments have proven that "Parity is NOT conserved" (unless, you also interchange matter and antimatter)
Now we return to facts that you WILL be tested on, on future exams. Spherical symmetry (for example, of a soap bubble) Axial symmetry (of a cone, unfertilized urchin or human oocyte, or an early urchin gastrula, or an early mammal blastocyst) One plane of mirror reflection symmetry (a pluteus, the exterior of the human body, The letters A, B, C, D, E, K, M, T, U, V, number 3) Two planes of mirror reflection symmetry (The letters H, I, number 8, 0)
Three or more planes of mirror reflection symmetry (A starfish, a sea urchin, a jelly fish, an individual coral animal, octopus, some other invertebrates, and plants;
diatoms are especially interesting examples, the symbol *, the letter Y, almost). Twofold rotation symmetry: (The letters N, S, Z, $, %, #, twobladed propellers for airplanes and boats) Threefold rotation symmetry: Threebladed propellers.
Glidereflection symmetry: WWWWW , approximately. Displacement symmetry: somites, ties along a railway, spots on an animal Legs on a centipede or millipede. "Displacementdistortion symmetry": ribs, vertebrae, legs on a crayfish.
"Dilation symmetry" Driesch’s discovery about embryonic regulation. "Dilationdisplacement symmetry" A long cone, or other shape that tapers in size. "Dilationrotation symmetry" A snail shell Remember Weyl’s definition: A thing is symmetrical, if there is something you can do to it, after which it looks the same as it did before. So if you have a pillowcase, or a sleeping bag (or some organ in the body, or some kind of invertebrate, that looks the same after you have turned it inside out, therefore it has turnedinsideout symmetry! (Which I just this minute invented!) Please try to invent some new kinds of symmetry.
Biologists have been left far behind: For example biologists usually consider only two forms of symmetry  "mirror image symmetry" (Plutei) and "radial symmetry", starfish. Radial symmetry is naively said to be evidence of evolution from sessile ancestors. Reduction of anatomical symmetry from axial to mirror image symmetry is said by biologist to be the development of symmetry.
Really, it’s the destruction of infinity minus one planes of reflection symmetry!
(NOT the formation of a new plane of reflection symmetry.) Despite backgrounds in physics and mathematics, Turing, Meinhardt and the advocates of "Clock and Wavefront" theories do not think of these mechanisms as breaking displacement symmetry. Liesegang rings also break displacement symmetry, and produce a displacementdilation symmetry (rings progressive farther apart and wider. Also slower to form?
Air pressure inside a soap bubble has spherical symmetry. In order to blow square bubbles, either the pressure or the tension will need to have less symmetry (the symmetries of a cube) The directional elongation of growing cartilages cannot be caused by stronger osmotic swelling in the long axis, because osmotic pressure is a scalar (and has spherical symmetry). The elongation must be caused by directionality of "Young’s Modulus" or possibly by directionality of tension.
Sperm entry location breaks radial symmetry of amphibian eggs
(stimulates formation of the anteriorposterior axis). In nematode embryos, Prof. Bob Goldstein proved that the location of sperm entry Also breaks symmetry, and stimulates formation of the anteriorposterior axis. In bird eggs, the direction of the anteriorposterior axis is believed to be initiated by the direction of gravity. In mammal eggs, it hasn’t yet been discovered what "breaks" either spherical or axial symmetry. But breaking of rightleft symmetry in humans is caused by flagella in the primitive node. This was discovered because of Karteganer’s syndrome, which is caused by a mutation that paralyses flagella and cilia. About 45% of people born with this syndrome have a reversed asymmetry of their aorta and stomach; another 45% have normal asymmetry of anatomy, and around 10% have partial reversal of right left asymmetry of internal organs.
