What is a Mosaic ?
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If you hear the term Mosaic, and your involved in the television industry, you might
think of an iconoscope which is the television camera's photosensitive surface.
Should you be involved in the Art World a picture or decorative design made by
setting small colored pieces of stone or tile into a surface might come to mind
as the meaning for mosaics. We see this on many floor and wall tile
designs. These are often attractive and colorful. Others may in vision the
process of making pictures or designs from a composite of overlapping
photographs as being a mosaic. In botany the term Mosaic is referred to as
a viral disease of plants, resulting in light and dark areas of the leaves.
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Biology use's the term mosaicism as a condition exhibited by an individual displaying tissues of genetically different types, or possessing cells of two or more different genetic constitutions or an organism that has two or more cell populations that differ in their genetic makeup. When a difference in color, pattern or physical structure exists for these two genetic makeup's we see a mosaic. Their cause could be environmental or genetic in nature but the end result is basically the same. They could either be the result of Somatic mutations or from Bipaterity sperm which is two sperms fertilizing a single egg. Regardless of their cause, different genetic cell make-ups result in different genetic production and when this results in something we can see with our eyes we have a mosaic. Lets start with Somatic Mutations. To find answers as they relates to our pigeons I contacted Dr. Lester Paul. Gibson editor of Pigeon Genetics, News, Views & Comments. With his permission I will cut and past portions of his emails regarding pigeon mosaics. |
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Somatic Mutations by Dr. Lester Paul Gibson Somatic mutations are common in plants. Many of these have been propagated by plant breeders and account for a lot of the plant variations on the market. Particularly of the plants that do not produce an abundance of seeds or that, produce seeds that are so diverse in their genetic traits that the desired trait is hard to get. Most of our fruit trees and many of our shade trees are propagated from grafts or cutting to keep the same genetic characteristics. These usually are propagated from somatic tissue. Some plants like Coleus are noted for their somatic mutants and easily propagated. I have seen somatic mutations on lots of plants that produce completely new varieties and thus somatic mutations are converted to new genetic status.
But that is another story since we normally cannot do that with animals. In animals, we see somatic mutations but they are usually not transmissible by tissue propagation. However, tissues from these can be grown in the proper medium. Somatic mutations include moles, warts, cancer, stray colored hairs, stray feathers, etc.. Mosaic areas can occur on varying amounts, sizes, and areas on animals just as they do in plants. Part of these can be somatic but most are apparently genetic in origin. The single feather or scattered small patches of feathers can be either genetic or somatic but by definition they are all mosaics. When we see what appears to be a somatic mutant, it many times turns out to be by selection, a genetic mutant. A couple instances of the latter readily come to mind: Way back when, a German family was rearing green finches from the Canary Islands, one bird turned up with a single yellow flight feather. By inbreeding, they were able to increase the yellow feathers until they had an all yellow bird, which is the ancestor of our yellow canaries. A number of years ago, I visited with Elsworth Kistler in Pennsylvania. He had an Archangel turn up with a white spot about the size of a silver dollar in the middle of its bronze chest. I saw the bird and tried to buy it from him. He wanted to breed more of the same and did not want to part with it. He did this but the white was now about half the bird. I purchased some of these from him. Breeding them together, he and I produced some all white with black tails. Breeding these together produced some all white birds. When these all white birds were bred back to full colored birds the white completely disappeared so it was a recessive white that acted as a migrational white just as the yellow was a migration mutant in the Canary.
Most somatic mosaics in our birds show up as a small spot or feather on
the body. If you have an ash red bird, a white bird, or an Almond with small
spots or feathers other than the color they ought to be (blue or black on red
ground) (red on blue ground) etc. they probably are somatic mosaics.
Many almonds will have a group of feathers forming a spot somewhere on the body and
these are probably somatic and not bipaternal. A somatic mutation can occurs in any cell (body cells) except the reproductive cells (genetic cells) i.e. genes on the chromosome which result in the reproductive cells (egg and sperm). In warm blooded animals it is fairly clear cut; unless the trait can be reproduced genetically it is considered to be somatic. In plants, this becomes blurred because somatic cells, as demonstrated above, can be selected to produce a new plant which can then produce its own reproductive cells and thus the somatic mutation becomes a genetic. In Insects and other cold blooded animals the action of mutations is blurred by the multitude of ways the organism can be, and is, affected by the environment in the very broad sense. Many insects can reproduce parthenogenically until the environmental conditions become stressful and then reproduce genetically. Many mollusks and annalids reproduce by being hermaphroditic. Some mollusks are born female and after a number of years change to males so that sexual reproduction can be accomplished. Some fish may change sex in the absence of available males in the population. Alligators and Crocs reproduce sexually but the sex of the hatchling is dependent upon the temperature of the eggs during incubation. All of these things have a bearing upon the picture as to whether a mutation in an organism can be called somatic or not. As you can see there is no simple answer. Bet you did not want to know all of this when you asked. Dr. Lester Paul Gibson |
Bipaterity sperm Mutations by Dr. Lester Paul Gibson I think in order to form a basis of understanding for the possible formation of what we call a mosaic pigeon; we must start at the cellular level. At one phase of the cell division, there occurs a different type of cell division from mitosis. This type of cell division is called REDUCTION DIVISION which forms the germ cells or gametes. This formation of gametes is known as maturation. We know that normally a new individual is started by the union of two gametes, a sperm cell and an egg cell. Sperm cells are formed by two cell divisions. Two primary spermatocyte chromosomes become paired closely in what is known as synapsis. Each splits in reduction division so that now there are four chromatids (a tetrad). These separate loosely and one from each dyad goes to each end of the cell and now the cells contain only 1 chromosome of each pair. The other two spermatocytes (the two not used above) divide by mitosis and form spermatids. These form sperm without further division. If during the reduction division part of each exchange sections with the other; we have what is known as a chiasma. THIS IS WHAT WE CALL A CROSSOVER. In the formation of the egg from potential egg cells, a similar series of events takes place. The only difference is that instead of the process producing 4 gametes (like the sperm production), three of the four are normally small and disintegrate. We call these POLAR BODIES. The fourth gets all of the cytoplasm and becomes a mature egg. Each of these cells will have one chromosome of each pair, the difference is that only one gets the cytoplasm and the others do not get any. If during this process instead of only one remaining, one of the others (a polar body) should also get some of the cytoplasm or the division is not complete and the polar body does not deteriorate; then we get a dual cell egg (a bipolar egg). In this case, each of the two poles may now become fertilized by a different sperm (bipaterity). Depending upon the size and viability of each; the area of mosaicism may be half or somewhat less than half of the resultant organism. Chromosomal aberrations occur in several ways but lets reserve our discussion to those that may produce mosaics. One type is called non-disjunction in which during cell division, both chromosomes go to the same end of the cell and thus producing a cell without a chromosome and one with both. This is not a theory because it is known to happen. Certain resultant combinations do not produce viable embryos and thus do not live and others instead of producing diploids may produce triploids, tetraploids, or monoploids for certain chromosomal pairs. Work with drosophila flies has proven that they can and do occur. Another type is called translocation (another crossover mechanism). Another type is called segmental interchange (a double crossover mechanism). Another type is called deletion (a mutational mosaic mechanism). It has been stated that a cell with three chromosomes cannot live. That has been patently proven to be false in many insect and plant genetic tests and manipulation results. It is known to even occur in a number of human cases in the sex chromosomes in which individuals may have the sex configurations of XXX, XXY, XXXY, XXYY, or even XXXXY. Somatic mosaics can occur anytime after fertilization of the egg. If early in the development, there may be near 50% of the organism or if later only a few cells. Let's consider the bipaterity sperm theory. If the sperm fertilize the egg at or nearly at the same time you would get a configuration somewhat like below: __ __ __ __ _________ Sperm A ------------------------------------------ Egg --- --- --- --- -------------------- Sperm B
As the egg starts dividing the Sperm A and the egg would produce cells
that become the right half and Sperm B and the egg would produce the left half.
I have the mounted bird. Dr. Hollander assured me that it was a
waste of money to have it mounted and preserved because everyone will just say
it is a hoax.
The Owl side is completely Owl with all the Owl attributes including the
pantaloons and the Roller side distinctly Roller with the distinct hetero
Baldhead marking. Even measurements of the beak show one side Owl and the other
Roller. Dr. Lester Paul Gibson
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The infamous two-sider
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As proof of the two sperm theory let me point to the existence of a bipaternal, bilateral 2 variety pigeon bred by Marvin Lee of Praire Grove, Ar.. This bird was a sooty, blue bar, baldhead roller on one side and a black homozygous Chinese owl on the other. A bilateral is formed at the first cell division or very closely to it forming one complete side different than the other. The fact that this specimen is of two different breeds, and involves more than just the sex chromosomes is all the more evidence of the two sperm theory. Dr. Gibson has the body of this bird preserved. |
What follows is an article written by Ken Davis. Ken is a close student of the late Dr. Willard F Hollander, a professor of genetics who first put forward the theory of two sperms fertilizing a single egg as one way to create a mosaic. Dr. Hollander specialized in the study of bird, mice and fruit fly genetics. He loved pigeons and was without a doubt the most knowledgeable expert on their genetic make up. The vast majority of what we know about pigeon genetics comes from him. Sadly he passed away and we can no longer ask him for guidance in our studies. So lets turn to Ken's article for more information. |
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Mosaic
Yes indeedy, that "two-sider" is a nice mosaic, for sure. But, something more. If you look at the photo's real close you'll see that the birds right side is a spread Chinese Owl, complete with the Frill, the Pantaloon, Owl shaped head, etc... Look at the head-on view.....notice the difference in feet structure, leg structure and length, the total package?
O.K., now,
we were all going over these pictures the other night at the Genetics Meeting at
Joe Fraziers Place, and Leon
Stephens told us that he knows how this happened; that it was "Bi-Paternity". He
said that the Frill and the Pantaloon on the Owl side of the bird are recessive
characteristics, so the Mother of this mosaic had to be a Chinese Owl, and was
bred by two cocks, a spread Chinese Owl cock and a blue series Birmingham
Roller, therefore the mosaic was Sired by two Fathers, hence the "Bi-Paternity".
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Well, this ain't what happened! The hen that
snuck into the cock pen was a "Brown Bar Birmingham Roller", and she was left in
there, or allowed to stay, and she was THE ONLY HEN IN THERE. Now, Frill and
Pantaloon being recessive characteristics, how'd they get there? How can they
express? Let's address the color part of it. The mosaic individual is spread
blue on it's right side, and blue bar on it's left side. Now, the Dam, or Mother
to this individual is a true Brown Bar, and she could only have been bred by
true Brown Bar Birmingham Roller cocks and spread blue Chinese Owl cocks. Well,
if she was "treaded" by both, ("Bi-Paternity"), the mosaic would have been a
Brown Bar Birmingham Roller on one side, and a spread blue Chinese Owl on the
other side. But it's not! Careful examination of this mosaic reveals that it is
phenotypically a "pure" Chinese Owl/spread blue on it's right side, and a
Chinese Owl/Birmingham Roller blue bar hybrid on it's left side. So how can this
be? The best explanation is "Super Numerary Sperm". That is to say that the
Brown Bar Birmingham Roller Dam was bred by a Spread Blue Chinese Owl cock, and
that more than one sperm from the Sire penetrated the ovum. And for reasons not
totally yet understood, the extra spermatozoa affected the phenotype to the
results which we now see.
PIGEONS IN GENETICS, FROM DARWIN
TO BIPATERNITY, By W.F.HOLLANDER,
Professor Emeritus of Genetics,
Iowa State University, Ames, Iowa50011.
This paper issued at the Department Zoology/Genetics, Iowa State University, February 2001. Not copyrighted |
My Summation
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OK that explains somatic mutation
mosaics and
bipaternal
mosaics. How are they
formed? Well I'm not 100% sure but here is my understanding as best I
can explain.
Only
one spermatozoid penetrates the egg membrane to fertilize it. That's
what’s supposed to happen; however, were discussing the abnormal not the
normal. Under normal conditions once a sperm has entered the egg a
chemical is released that kills or at least prevents the remaining sperm from
penetration. Some hens are deficient in the production or release of
this chemical and are therefore prone to production of mosaics. Thus two
sperms on occasion do enter the membrane before the defensive mechanisms can take
effect. When this happens
and a
bipolar egg exists then each pole may get fertilized by a different sperm (like
our two-sider) or; in
the case of an egg that is a normal polar body,
chromosomes from one spermatozoid pairs with the matching chromosomes of the
egg while the remaining spermatozoid becomes surplus DNA within this newly
formed first cell. As the mitosis process for body development and
growth continues this surplus DNA either replicates and is passed along as
more surplus material in the new daughter cell or they don't replicate and the
new cell is free of it and the potential for a mosaic is lost. Should
any of this surplus DNA replicate there is a chance for a
crossover mechanism to occur
in the chromosome chain of the new daughter cell (the other half of the
divide) and a mosaic is formed. As the embryo grows it continues
to develop and divide with this new string. The earlier an occurrence of DNA swap happens the larger the mosaic patch will be.
The numbers of sperm cells entering an egg does not change its sexual outcome.
The sex of any pigeon, mosaic or not, is determined by the hen's egg not the sire's
sperm.
When the hen’s egg comes with a Z chromosome you get a cock Z//Z. If
she donates a W chromosome you have another hen Z//W. The embryo
will grow from that first single cell. As it replicates it becomes
two and then four and so on number of cells. Any extra DNA from a surplus
sperm is either disposed of or it displaces some of the original DNA from the like chromosomes united in that cell. For base color
it would involve the Z chromosome but it could involve any of the 40 some
chromosomes sets. This replication process is going
on for each body cell. Should surplus DNA displace the original then that cell and all future cells produced from it would continue to
contain this new DNA combination; assuming of course that the remaining DNA is
then discarded.
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So how does that effect reproduction for structure and color of the offspring? Since the genetic markup of the cells located in the gonads determines the DNA provided to the egg or sperm, the cells which make up these gonads will be the DNA mix that the bird will reproduce, regardless of what the remainder of flesh and feathers are genetically.
Here's another mosaic but with a very subtle effect. It comes by way of Layne Gardner. This is what Layne had to say about the bird. "I am guessing that right side (bird's left wing) of this clean leg Field Pigeon is expressing some influence of the ice factor. The left side is common blue white bar. As we examine our birds for these more subtle mosaic effects it seems apparent that mosaics may appear more frequently than once supposed." Layne Gardner Colton, California
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A mosaic of Mosaic photos

Five Mosaic photos by Michael Spadoni of Geelong Australia
Half Smoky / Blue bar by AL Neese and a Half Mealy / Grizzle by Larry Davis
These two photos of the
same bird were taken by David M.
Longseth at the 2003 NYBS.
The right wing shield is a blue check while the left side is a dark 'dun/brown' color with heavy black flecks or sootiness.
This left side may be almond as the sire was an almond bird.
This young homer cock belonged to Timothy Hume of Saltspring
Island, BC Canada.
It was killed at age four by a falcon. The bird was both a blue
bar and a mealy.
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