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A Discussion of the Micariff Phenotype; misconceptions and facts PDF

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MONDAY, MARCH 30, 2015 A Discussion of the Micariff Phenotype; misconceptions and facts © Alan S. Bias Permission granted for nonprofit reproduction or duplication of photos and text with proper credit for learning purposes only. March 30, 2015 March 31, 2015 (Updated) Blond Micariff, courtesy Micke Norberg Grey Micariff, courtesy Micke Norberg According to California breeder and IFGA member Mike Khalid, the Micariff phenotype was created in Sri Lanka by two breeders from his homeland, Michael Cole and Denis Ariff. The name itself a truncation of Michael's given name and Denis' surname: MicAriff. While Micariff is most often associated with a reduced melanophore blond (b) phenotype, it is also recognizable in wild-type grey with increased melanophores. As a pedigree stock breeder I am hesitant to assign specific genes to produce expected result (phenotypes). If for no other reason modern research and breeding’s have shown the many phenotypes are not the product of “single genes”, rather the product of “multiple genes” or mutations upon them in combination or linked complexes. To this reasoning, as in all my writings, will refer to a gene(s) expression as a trait. Page 1 of 21 The required traits needed to produce the Micariff phenotype have long been the subject of conjecture based on both purebred and outcrosses maintained by breeders. Results from each suffering from lack of scientific method needed over multiple generations to provide proper ratio’s for analysis. Yet these breeding’s can still tell us much from visual records. The two most prevalent theories having arisen in Europe and Asia where Micariff’s have a stronger following by serious breeders in comparison to North America. The Asian version while lessor defined basically states that the end result is the product of a metal trait, likely Metal Gold (Mg) + Snakeskin Body (Ssb) + Snakeskin Tail (Sst). [Note: Hereafter Ssb / Sstwill be referenced singularly as Ssb]. The better defined European version asserts a product of homozygous Stoerzbach1 (s) + homozygous snakeskin (Ssb): Males = XSsb YSsb ss mgmg and females = XSsb XSsb ss mgmg. It should be taken into account here that many European breeders do not make a genetic distinction between Stoerzbach and several other metal traits as is common in Asia and North America. Grey Micariff female, courtesy Myster Green Blond Micariff female, courtesy Raj Seshagiri Hypothesis: In response to both of these theories, I state each as being only partially correct. I propose: That Metal Gold (Mg), Stoerzbach (s), Snakeskin Body (Ssb) all in homozygous state, and a “missing trait” are required to produce the original metallic Micariff phenotype in full body expression. That this missing trait is required in conjunction with Steorzbach a "full body" trait acting as a precursor to effect the "metallic gold Micariff Page 2 of 21 phenotype" (hereafter metallic) in both body and finnage. As a result two actual expressions are commonly produced when Stoerzbach is lacking or in heterozygous form; metallic and non-metallic body (hereafter non- metallic). [Note: It should be stated that in a true breeding metallic strain females under this hypothesis would need a genotypeXSsb XSsb ss mgmg in females and XSsb YSsb ss mgmg in males. For for producing a percentage of metallic a genotype heterozgous for stoerzbach in either males or females would suffice. For production of non-metallic a genotype lacking Sterozbach in one sex or heterozgous for stoerzbach in both sexes would suffice.] Homozygous metallic; XSsb YSsb ss mgmg, courtesy Mitchell Weintraub Heterozygous metallic offspring; X YSsb Ss mgmg, courtesy Mitchell Weintraub Heterozygous non-metallic; XSsb YSsb Ss mgmg, courtesy P. Shaddock Discussion: When discussing genetic makeup and expression of color & pattern in guppies, one should undertake an understanding of what we are actually seeing with the naked eye. You quickly come to the realization we are not looking at a colored fish, but rather it's reflective qualities. The Bagnara Dermal Chromatophore Unit was first published in 1968. For nearly four decades it was for the most part ignored by both the scientific community and that of the Domestic Guppy breeder. Though the latter Page 3 of 21 often stumbled upon it in breeding's and produced visible results without understanding the genetic implications. Longtime successful stock breeders breed by eye; through the power of observation to achieve if not understand results. We are visual thinkers to varying degrees. Little different from a musician who plays by ear. This shared trait among breeders has met stiff resistance in acceptance by many. Yet even the scientific community has slowly started to support what we as breeders have known for centuries; that successful breeders have an inherent and often above average ability to observe the natural world and create domestic results (Arnheim 1969 & Grasseni 2004 & 2005). Our shortfall in the eyes of science has been an inability to communicate this understanding to others, both outside and among our peers. In the following simple visual aid, used in support of Bagnara's work on color study, we have the basis of his theory as published in multiple papers over a period of two decades. [Note: color added to black and white photo to reflect chromatophores.] The Dermal Chromatophore Unit; Bagnara (1968) & Bagnara et. al. (1973), reprint from Grether (2004) [Excerpt Grether (2004): The dermal chromatophore unit... ...of a hypothetical green frog, showing how the xanthophore, iridophore, melanophore and underlying fascia layers interact to determine the overall colour of the animal. Wavy lines depict the paths of light of differing wavelengths through the cell layers...] An understanding of the Dermal Chromatophore Unit in conjunction with an understanding of genetic associations between melanophores - color pigments - iridophores has much to offer serious breeders. The two will help plan your mating's and alleviate trial and error through random breeding's. In the end help understand your results, both positive and negative. Even though your goals or that of a show standard may call for a specified result, is it in reality genetically possible? Tank space is too precious to "recreate the wheel" in re- producing the failures of our predecessors. Yellow color pigment is often very motile and hard to define by the naked eye unless put under a microscope to determine composition. From a breeders perspective I tend to make a couple basic assumptions in associating yellow traits when it comes to yellow color pigment cells. Page 4 of 21 The assumption is based on perceived mode of inheritance and motility. If autosomal dominant or recessive, likely metal gold. If sex-linked, likely yellow color pigment. Even this is not guaranteed. There are so few identified “yellow color pigment” traits in the body. If there is an “expected norm” it might be: 1. Yellow color cells are often very small and round. 2. Red color cells are larger and less defined. 3. Wild-type orange seem to fall in between in both size and shape. Working from memory, wild-type yellow cells are most often punctate; small & round. They can also be dendritic; large with arm-like extensions. The latter can result in regions with dark dense and motile yellow coloration. Population type (punctate or dendritic) and numbers often differ depending upon a lower layer of leucophores or iridophores in both body and finnage. When high concentrations of either leuchophores or iridophores appear absent to the naked eye (clear finnage) dendritic & punctate are often found together close to the body. As you extend further out on finnage dendritic reduce in numbers, while punctate continue. Even then, this is subject to both type and concentration of melanophores. Both red / yellow cell placement and population is in interaction with melanophore types and location. Both punctate and dendritic yellow color cells are found in various locations in body and finnage. Punctate throughout the body and dendritic concentrated in specific locations. Based on breeding results, trait definitions, and an understanding of trait composition it seems dendritic are the most motile. Punctate are scattered about having less direct interaction with melanophores. Dendritic tend to collect around melanophores. The primary difference often determined by leucophores (white), iridophores (silver / blue) or neither underlay color pigment cells. [Note: Motility of yellow color cell pigments over iridophores or leucophores allows for increased expression by positive selection.] The two best examples for understanding yellow color cell population and color type are Schimmelpennig Platinum and Full Platinum strains. Schim Plat is a “fixed” trait, likely in complex. Yellow coloration in Schim Plat is comprised of evenly dispersed punctate cells and areas of collected dendritic cells. Line of decent plays a large role in Schim Plat composition. Schim Plat is a full body trait derived from Vienna Emerald Green. Vienna Emerald Green in turn derived from “wild-type”. Wild-type is comprised of both silver / blue iridophores and Le in well-defined zones of regulation. While the Schim Plat has distinct zones of Le regulation, it is overall a silver / blue iridophore full body trait. [Note: Schim Plat in outcross often expresses reduced levels of Mg yellow coloration in F offspring. This in itself is suggestive that we should view Mg not as a “yellow / gold 1 iridophore”, but rather as a “yellow / gold color pigment trait”. There is limited evidence pointing to the existence of either red or yellow iridophores.] Full Plat (NiII + Le) is the product of combination, easily segregated in outcross. The increased expression of Le results from enhancement through combination with a full body trait; NiII. Yellow coloration iin much of the body is predominantly comprised of evenly dispersed punctate cells over white. Iridophore regulation, while present, is minimal. In IFGA circles the Micariff does not command the respect of many breeders as having what is needed to produce a fish worthy of competition in show. Many breeders have attempted to do so, with marginal results, Page 5 of 21 and in the end abandoned the process in favor of more reliable traits to produce a Yellow Guppy. Others have incorporated selected traits into existing strains with positive result. Melanophore reduction will result in increased visibility of color pigmentation; i.e. Blond Yellow or Blond Full Gold appear more yellow than grey counterparts. However, without a corresponding increase in reflective qualities, color may become flatter than grey counterparts. The blond Micariff Guppy is a phenotype that is either loved or hated by individual breeders. It has contained in its genotype in homozygous form the traits needed to produce a Yellow Guppy. Although this is often with corresponding reduction in overall finnage. An outcross can result in near complete loss of yellow color pigment or Mg if you lack an understanding of the genetic composition. The traits most often favored in production of yellow strains involve combinations of metallic iridophores; Metal Gold (Mg) or Schimmelpennig Platinum (Sc), melanophore reduction or masking; Blond (b) or Albino (a), metal enhancement; Stoerzbach (s) or Full Gold (NiII + Mg), and red color pigment removal; Asian Blau (Ab). Due to the very nature of yellow color pigment interaction with melanophores and iridophores, results while impressive have fallen short of achieving a full Yellow Guppy. By some considered genetically impossible and likely so without the further identification of traits in the future. While Stoerzbach and Snakeskin are both considered full body traits, it is long known by breeders that neither the combinations of ss + Ssb or Mg + Ssb will in themselves produce a “true” Micariff phenotype. At best each will produce a “co-expression” of either combination not effecting a full body metallic Micariff expression. Numerous breeders are aware that F offspring from either sex Micariff parents in reciprocal outcross will 1 produce a snakeskin phenotype. This in itself is suggestive of Ssb in either sex being in a homozygous state. In turn, sib-breeding of F stocks will again recombine in subsequent generations to produce Micariff’s with 1 increasing numbers. Here in outcross and re-combination of s and Ssb is where we first see clear evidence of our missing trait. In various Micariff phenotypes, you will notice distinct variations in color and pattern. Some are more metallic in nature, while others have higher expression of "flatter" colored leucophores. It is common for autosomal traits to produce partial expression in heterozygous fashion. With high expression of leucophores vs. iridophores a red shoulder stripe is often apparent, especially in blond males. This is often referred to as an "Old Fashioned" shoulder stripe; a by-product of Vienna Emerald influence. When faint lavender colored the result of Purple Body Mutation (Pb). When red, the result of X-link red color pigment. In all cases, there is a direct correlation between increased anterior iridophores and a reduction or complete masking of the shoulder stripe. Page 6 of 21 Outcross has led to the further development of several other Micariff phenotypes over the last couple decades by both Domestic Guppy Breeders and Commercial Farms. Under the strain names of; Berlin Buttercups, California Golds, German Yellows, Tequila Sunrise, Sunset Guppy, etc… Each the result of various states of zygosity involving the original foundation traits &/or addition of new traits to include; Half Black (NiII), X- link red color pigment in finnage, Lazuli (Lz), and Japan Blue (Ca)2. Sunset’s are the simple product of the inclusion of X-link red color pigment in heterozygous or homozygous fashion, as can be demonstrated by breeding results on a Punnett square. They are not the product of a distinct “Sunset Micariff” allele as has been proposed & promoted by some breeders. Red color pigment was initially infused as a heterozygous X-linked gene. As result of crossover red color pigment can, and often of does, become Y-link. Expression in Sunset’s will vary between heterozygous and homozygous state. Where is our evidence of a full body trait and zygosity impacting expression of red & yellow color pigment in a Sunset Micariff? It lies in the pattern expressed by the two chromatophores; a clear delineation in co- expression of the red (erythrophores) and yellow xanthophores). In most instances yellow color pigment in guppies, depending upon linkage and zygosity, is epistatic to red. The end product being either yellow or orange (wild-type orange). In the Sunset Micariff, while there may be minimal blending of the two color pigments, the expected result is clear delineation; a repulsion of red away from yellow to the exterior edges of finnage. To produce such a result, a “full body” trait is required. In example: NiII produces a similar repulsion between yellow color pigment and black melanophores in finnage. Thus, demonstrating its effect as a full body trait. Page 7 of 21 Y-link Half Black (NiII), courtesy A. S. Bias In further example of pigment repulsion, we have the new “Sunburst Swallow” phenotype of Asian Breeder Akrawat. While rather fluid in makeup and expression, it relies on co-expression of NiII + red & yellow color pigments. Though he often adds Schimmelpennig Platinum (Sc)3and Ssb traits to produce further variation. As in the Sunset Micariff, while there may be minimal blending of the two color pigments, the expected result is clear delineation; a repulsion of red color pigment and black melanophores away from yellow to the exterior edges of finnage. In Sunbrust Swallow the full body trait is NiII or Moscow Blau Additional Gene (MBAG). [Note: The presence "swallow" trait and not only it's effect on finnage, but also confinement of yellow color pigment between hemirays.] Sunburst Swallow showing red pigment repulsion in both males and females, courtesy Akrawat Farms. Now we are to a point where it is prudent to reveal the missing trait found in Micariff I have been alluding to. Leucophore White (Le) has been around for a number of years, though seemly overlooked by many. Yet is found in genotype of many solid colored strains and frowned upon for hindering additional color pigment coverage. In example the following photos with minimal expression similar to wild-type. Page 8 of 21 Albino Full Red, courtesy of Paulo H. Keijock Muiz Expanded expression of Le has been known by long-time IFGA breeders as "Parrish White" in the anterior shoulders. In his writing's Phil Shaddock refers to as leucophore body (Le) and leucophore fins (Lef). While independent regulation between body and finnage has long been accepted in general, I am no longer sure this is a valid assumption in general. For some time I have been referring to this trait as “Micariff White” (hereafter Leucophore White) in the body while in correspondence with fellow breeders in discussing the Micariff strain. Here is where I will step out on a limb and hope it does not break. To my knowledge not a single "solid" Le body and finnage strain has ever been created that does not also incorporate at least one sex-linked or autosomal "full body" trait. [Note: White guppies are normally a combination of; 1. NiII + Le. 2. American Pink White (Pw) and one or more or the following Ivory (Iv) or Asian Blau (Ab) or Sc or s]. Rather than devote extensive space to speculation and further explanation to Le qualities will simply state; Leucophore White pattern is likely the result of genetic switching in utero. Possibly it is a form of conversion between melanophores and leucophores. As can be seen in the following IFGA examples, modification of Le often appears limited to expression in the anterior body. Each of the three males lacks a visible full body trait. IFGA breeders have only been marginally successful in masking anterior Le with inclusion of iridophores by Viridis (Vir), i.e. "blue belly" trait, Full Red body traits or Mg to produce a green cast in the anterior belly / shoulder region. Page 9 of 21 (clockwiset) IFGA Blue, Purple and Green deltas, courtesy of Bryan Chin. In this example an IFGA Gold Delta expresses finnage comprised solely of Le, less addition of Mg and Variegation (Var). It is a true solid white fin strain encompassing at least two full body traits; b and Pb. Leucophore White is present anteriorly, incompletely masked by Mg and Pb. I like to view Mg as an opportunist; It randomly seeks out regions lacking color pigment from fixed traits. It is amplified by full body traits, to populate large areas in regions comprised of iridophores or modified leucophores. Gold Delta, courtesy Bryan Chin In this example a Half Black White (IFGA HB AOC) phenotype is comprised solely of Le in finnage. Or is it limited to finnage? It is a true solid white fin strain encompassing a full body trait; NiII. While white coloration of scales on the topline is often simply equated to "platina", it is the result of Le in co-expression with NiII. Platina is easily modified to red, yellow or blue. Page 10 of 21

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