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Monday, August 13, 2012

Ginga Sulphureus... ...Evolution of a new strain or variant upon existent?


© Alan S. Bias
Permission granted for nonprofit reproduction or duplication of photos and text with proper credit for learning purposes only.

August 9, 2012

Ginga Sulphureus   ...Evolution of a new strain or 

                                                      variant upon existent?

F2 Ginga Sulphureus


Introduction:
It seems most new strains come about from one of four scenarios:  1. Random hybridization. 2. Mutation.  3. Crossover.  4. Planned modification.  The latter should stem from a predetermined breed plan with well set goals working with a recognizable parental strain(s) and any additional genotype you wish to infuse.  In such instance one of the first questions that comes to mind is just how much modification is needed for distinction as a new strain from it's parent?  When is it a new strain and when it is just a variant of the parental phenotype?


Discussion:
So what do we consider a variant?  In a simple sense when limited sex-link or autosomal traits are introduced.  Such as blond, albino, another body or fin color.  Many breeders will often limit their discussion of variants to autosomal effect. In a strict sense whenever a breeder acquires a set of genetics, maintains it for several generations in his/her own fishroom under their own devices it becomes your strain.  Even when bred towards original well defined standards of the creator or a breed standard.  This results from a change in both environmental and genetic inputs that, while similar, are no longer under the same controls.



Ginga Rubra - photo K. Tanaka with permission
A couple of years ago I first saw a picture of Kenjiro Tanaka's Ginga Rubra.  Ken has been well known in Killifish circles for many years.  His 30 plus years of manipulating guppy genetics is far less known to breeders outside of his native Japan.  In email correspondence and review of his website:  http://www.ne.jp/asahi/medaka-ken/k.t/index.html it becomes clear Tanaka's interest in guppy breeding favors non delta strains that are small, highly active, with alot of coloration.  In variations of his Ginga you can find finnage that parallels caudal shape found in some killifish species.

Ginga Rubra - photo K. Tanaka with permission
For quite some time I pondered the Ginga Rubra phenotype and whether it would hold my interest.  A first step of any breeder when considering acquisition of new genetics should be an analysis of the potential makeup and it's value to your current breedings.   It was  apparent that a fair amount of the Ginga traits were autosomal and that the phenotype would always be fairly fluid in many respects.  Similar to how I maintain most of my Swordtails.  To those who appreciate non-delta type guppies Ginga are quite stunning to the eye, but in the end still a red dorsal guppy and they rarely hold my interest for long.  Which I have always found strange in that my primary focus in large animal livestock breeding has always been towards red:  Red Galloway cattle, Moorit Shetland sheep, Rhode Island Red chickens, and more.  So, after looking over limited photos of the strain for a number a months I felt I had a decent understanding of the genotype comprising this strain.  The honor of defining Ginga Rubra as a strain belongs to Tanaka as the breeder. In correspondence he indicates, "My Ginga strain is extremely variable and I'm working with many types to get (it) established."  I'll only list, describe and then discuss several visible components.

Ginga Rubra - photo K. Tanaka with permission
Today most informed breeders are inclined to view genotype based on gene complexes governed by regulation, rather than single genes resulting in specifics traits. The Overall Ginga phenotype is comprised in such a manner.  Where it differs from many modern strains is being a complex composition of recognizable traits described many years ago on a basic wild-type form.  Ginga genotype is not hidden like many modern strains if you know how to look.  You might describe it as a series of overlays and overlap.  Very similar to the Vienna type Swordtails I have worked with for nearly 30 years.

In total the Tanaka Ginga Rubra phenotype is bred for and contains at least the following visible traits:  Blond (IFGA gold), wild-type, possibly reduced Emerald Green Iridescence (EGI) from no visible peduncle spot, Iridescence, purple body mutation, barring overlayed in purple, roundtail (though some males exhibit sword-like extension), ocular spotting (in peduncle base into caudal), increased number of Wingean "orange spots" along the lateral line, red dorsal (sex linked), multiple genes identified by Winge and Kirpichnikov for caudal color patterns and shape.  

Winge, O. (1922b), male exhibiting Sulphureus and Rubra

O. Winge in his 1922b and 1923 papers describes both the sulphureus (su) and ruber (ru) genes and hints at the possibilities of crossover to explain phenotypes with only partial components of each.    In Winge's 1927 publication he goes one step further and not only illustrates this process of crossover, but reduces both su and ru from two complex genes into to four distinct genes.   The complex sulphureus becomes vitellinus (vi) and coccineus (co), while ruber becomes minutus (mi) and sanguineus (sa).  Why is this important?  Because it forms the basis of multiple components of Ginga Rubra and Ginga Sulphureus.  In 1922b he describes each:  Sulphureus;  1.  Sulphur yellow colour in the dorsal fin and a dark dot that only at times is visisble in the said fin.  2.  Sulphur yellow colour in the tial and in the caudal fins.  3.  Red colour in the lower edge of the caudal fin.  Rubra;  1.  Red colour proximally in the upper edge of the caudal fin.  2.  Large oblong red side spot, lying for the most part below and behind the dorsal fin.  3.  Dark side-dot in the tail at the base of the caudal fin.


Winge, O. (1922b)

Winge, O. (1927)


Winge, O. (1927)
Winge further describes in his 1927 publication two other genes that are potentials in the makeup of Ginga.  Those being Tigrinus (Ti); an X-link consisting of 1-2 black pigment stripes, and Zebrinus (Ze); a recessive autosome consisting of 2-5, generally 3 dark pigment stripes.  He indicates they resemble Ti, but are more pronounced.    As an autosome this trait can be both heterozygous or homozygous in form.  A heterozygous sire will statistically pass the trait to 1/2 of his sons, but not necessarily expression.  This is determined by the recessive or dominant nature of the autosome.

Winge, O. (1927)




Phang, V (1999)
There is a 3rd form of barring described by V. Phang, et al. (1999).  In conclusion she proposes:  "The designation of (bar) for an autosomal recessive gene, when in homozygous condition barbar and present with snakeskin body pattern (SSB) produces a vertical barred pattern".  The dominant and allele being bar+.  There still remains alot of speculation among breeders if bar and Ze are not one in the same.  Source fish listed were limited to a strain of Y-linked Singapore Yellow Snakeskins.  No where in Phang (1999) is there any reference to reciprocal outcross of bar+ Y to non snakeskin strains composed of barbar XX or barbar+ XX females.  Therefore, it is possible that bar and Ze are one in the same as her statistical analysis does not prove bar and SSB were found in complex.  Simply co-expressed on males as individual traits.

Iridescens (ir)  Winge 1922b, 1.  A characteristic mother of pearl sheen on the body.  2.  2-3 red smaller side spots.  3.  Black side-dot on the tial near the caudal fin.  4.  Black side-dot on the anterior body.

Winge, O. (1922b)
Ferrugineus (fe) Winge 1922b, 1.  Black rust-coloured part in the proximal part of the cuadal fin.  2.  Black side-dot in in the tail near the caudal find (viz. with ocular spotting in the caudal).

Winge, O. (1922b)
Purple Body Mutation - undescribed, is a very important Ginga trait that needs to me mentioned here.  Tanaka's Ginga Rubra are bred exclusively for Pb.  To the best of my knowledge there is no formal description of this trait in guppies, having found no published research or reliable documentation of it.  Discussion with other breeders and researchers has also provided minimal insight.  To my knowledge this trait is found in only two of three published members of the guppy family comprising the subgenus Acanthophacelus; Poecilia reticualata, Poecilia obscura, and Poecilia Wingei.  This trait does not seem to occur in any of the pure Poecilia Wingei (Enders) that I worked with, nor are there any photos online from reliably documented populations.  It is only expressed in hybrids.  In the past, having been able to breed for it reliably, it did not seem necessary to set up breeding groups, dedicate tanks and document results statistically.  My test crosses to decipher it, in several strains, to include Ginga, Vienna and Wild-type, are not yet conclusive.  

This trait I refer to as Purple Body Mutation (Pb).  It's counterpart is Green.  My test breedings to date with non Ginga strains point towards Pb being an autosomal recessive trait.  Green presumably could be a dominant wild-type allele or epistatic trait as evidenced by lack of Pb in Endlers.  Until it is proven whether the latter is a wild-type allele (Pb+) or a distinct trait (Gb) I see no use in defining it as either.  To do so would require a larger number of free tanks than I have available.  Two strains; one homzygous for purple and another for green.  Documented analysis of reciprocal F1 crosses followed by F2 sib breedings.   
Vienna LS in both Pb (upper) and green (lower)
Purple Body Mutation as an autosomal recessive trait is a precursor in determining visible color resulting from pigment overlay.   Pb is what allows for creation of a true purple phenotype (viz. IFGA Purple).  Effect is limited to certain regions of the body.  Always in the peduncle in strains with reduced iridophores in this area.  In conversion of Wingean spots from orange to pink.  The expression can also extend along the shoulders and into the topline.  In most males, the belly region does not appear to be altered by the effect of Pb.  Green males express Wingean spots in orange.  Pb males express Wingean spots in pink.   While Pb is most noticeable in the peduncle region of male guppies, it is also expressed in females.  Along the lateral line and in the shoulders towards the topline, appearing as an iridescent purple sheen on the scales.  


Breedings:
In late summer of 2011 I arranged to obtain a trio of pure F1 Ginga Rubra from an individual who had imported directly from Tanaka.  My only requests were a male having a round tail and ocular spotting with no visible extensions, roundtail being a recessive not easily identifiable in my Vienna.  This request was fulfilled with the above below as he appeared after 4-5 months growth.
Ginga Rubra (P sire partially colored)
Ginga Rubra (P sire mature)
 As some of you have surmised based on yellow based breeding program my primary focus, at least initially, would be to remove red  in the dorsal and caudal via selective breeding.  If not feasible to at least mask it via epistasis from traits found in my Vienna LS.  To do this I would initiate breedings with a pure Ginga male and several pure Vienna females, blond and grey, to include both green and purple body mutation.  Then utilizing F1, F2 and F3 hybrid males exclusively bred the first 2-3 generations in backcross to pure Vienna females.  All hybrid filial sibling females were culled prior to onset of maturity.   Such a breed plan would avoid the pitfalls of segregation when using siblings.  In result:  (((Ginga * Vienna LS) * Vienna LS) * Vienna LS).  All initial Vienna females came from my Line1 as this line does not posses partially dominant autosomal Z-bar.  It does contain both Pb and green in composition.

One of the first things I wished to know about about Ginga had to do with barring.  Was this trait traditional autosomal Z-bar or the elusive recessive sex linked Tigrinus as documented by O. Winge early in the last century.

Ginga Sulphureus (F1 sire)
The F1 surprisingly resulted in at least two distinct phenotypes from several litters.  One was very Ginga in appearance to the sire with the addition of yellow finnage as hoped.  From the available males the one above was selected to sire the next generation.  You will notice a residual amount of red in the upper caudal.  While I did not do test crosses, this likely was the result of Y-link mi as it does not manifest in my Line1 Vienna stocks.  Dorsal size and shape was near identical between father and son at maturity.  Moderate in length and tapering to a well defined point.  Retention of overall body pattern still hinted at Y-link &/or autosomal traits.  In both F1 phenotypes barring was present in 100% of males from litters of four (4) females to varying degress  This without a doubt says autosomal Z-bar and not sex-linked Tirgrinus.  It also suggests an autosomal dominant version of Ze and not a recessive version as my Line1 Vienna LS traditionally lack Ze.  In F2 backcross Ze is still present nearly 100%, albeit weakened in some males.  This continues to suggest that the Ginga are comprised of a autosomal dominant version of Ze and not a recessive.  You will notice the appearance of a green peduncle spot in this F1 male.  It would be hopeful to suggest an autosomal input from his Vienna dam.  It is more likely visible from a reduction in barring.  Z-bar is often epistatic to peduncle spots when bred for posterior coverage toward the caudal base.  

F1 Ginga * Vienna hybrid
I mentioned a second distinct F1 phenotype that appeared in both blond & grey siblings.  A single grey female likely was the dam of all.   I have gradually culled or given away the blonds, but kept back a lone grey male who I am now breeding to both pure Vienna and F2 Ginga Sulphureus females.  His finnage is very reminiscent to the types of guppies I bred as a youngster back in the late 1960's and early 1970's.  An interesting Lyretail with little extension and in appearance mimics many Aphyosemion killifish species.  It will be interesting to see how much of this caudal type was passed, from his pure Vienna dam.  I have always suspected that much of either Y or X-link DS trait is carried by females both in sex link and autsomal, and is masked by epistatic Y-link LS.

F1 Ginga * Vienna hybrid
Outside of his caudal fin, other body color / pattern components are still very Ginga.  Note that all F1 males of this phenotype lacked dorsal color pigment.  Either being  clear or lightly colored  by metallic (Mg) and this is probably in complex with the unique caudal shape.  If so, a possible Y-link as my females will pass X-link yellow dorsal nearly 100% when linebred.  These males also lack a green peduncle spot.  In both F1 phenotypes the amount of orange / pink spotting seemed to increase and intensify with age.  Both in anterior regions and over posterior barring.

Segregation in the F2 has come about in the expected ratios.  While Ze has remained strong, the degree of intensity (viz. total number of bars) has decreased in some males from several generations of backcross.  A mixture of blond / grey in both green and purple body mutation.  For F3 breeding groups of each have been set up with sibling females to further this new strain at least one more generation.  With emphasis on increasing what I am sure Kenjiro Tanaka would consider not only a flaw, but one of his greatest challenges in breeding this unique strain.  Removing or at least masking the peduncle spot while increasing both ocular caudal coverage, also known as "peacock spots", and the degree / intensity of Wingean orange / pink spotting.



F2 Ginga Sulphureus
F2 Ginga Sulphureus



F2 Ginga Sulphureus



F2 Ginga Sulphureus
F2 Ginga Sulphureus













F2 Ginga Sulphureus

Conclusions:
I come across quite a few comments in correspondence or on the web hinting at Ginga's deriving much of their phenotype from possible Poecilia Wingei (Endlers Livebearer) influence.  Surprisingly, all of the individual traits I identified, less some finnage, were already existent in either the Line1 / Line2 or both of my combined Vienna Emerald Lowersword (LS) strain.  Those few that are not common today, such as ocular spotting, were often found in guppy strains of the past.  This has allowed me to utilize Vienna in conjunction with a new direction in breeding Ginga's.  That being a reduction of red finnage in favor of pure yellow.  The primary trait in found in Ginga Rubra lacking in my Vienna LS is the layering of purple striping over Ze.  I also found that one of the most striking traits on the Ginga to be a seeming ability to express both Pb and green simultaneously.  Again, this trait while rare is also found in P. Reticulata strains.   The male on the right has no Ginga genetics in his makeup, yet posses the same ability to express both orange anterior and pink posterior.  Note that I have never run across an inversion of this combination, pink in the anterior and orange in the posterior.

F1 Ginga Sulphureus (l) and Schimmelpennig Platinum DS crossover male (r).
 Both expressing Pb posterior in peduncle and orange anterior in shoulders.
Breedings continue to indicated Pb as a recessive autosome.  Nothing points toward it being (partially) dominant as in the case of Asian Blau (Ab).  Analysis of several prior non Ginga breeding test results, consisting of Vienna / Schim / JB albino, and lack of Pb in Endlers point to green being a wild-type allele Pb+.  As Tanaka line breeds for Pb in Ginga Rubra, it is assumed both males and females are homozygous for the Pb trait.  Yet, no green presented in F1 from a pure Ginga Rubra single sire mating to 2 females (Ginga * Vienna LS).  It is doubtful both of these females were homozygous for Pb as their sire had been green.

I'm looking forward to a series of reciprocal Vienna LS * Ginga Sulphureus crosses a couple generations from now.  It will be interesting to see what autosomal traits will be contained in the genotype of linebred  Ginga Sulphureus females.  In the long-run when I work with a new strain the end result is not only to see what traits I can stabilize, but identify those which I can feasibly incorporate into my Vienna LS.


References:
Winge, O., Genetic Laboratory of the Royal Veterinary and Agricultural College, Copenhaen
(1922b).  "ONE-SIDED MASCULINE AND SEX-LINKED INHERITANCE IN LEBISTES RETICULATUS"
(1923).  "CROSSING-OVER BETWEEN THE X- AND THE Y-CHROMOSOME IN LEBISTES"
(1927).  "THE LOCATION OF EIGTHEEN GENES IN LEBISTES RETICULATUS"

Phang, V. et al., Dept. of Biological Sciences, Faculty of Science, Nat. Univ. of Singapore (1999). "Interaction between the Autosomal Recessive bar Gene and the Y-linked Snakeskin Body (Ssb) Pattern Gene in the Guppy, Poecilia reticulata"

Tanaka, K & Bias, A. S. (2012).  Email correspondences on Ginga Rubra



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Which do you find of more challenge as a breeder, creation and stabilization of a new phenotype or continued propagation and improvement of and existing one?
  Each a unique challenge in their own right.   

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3 comments:

  1. At your conclusion you say...."continued prorogation and improvement....." Prorogation is the dissolution of parliament as we have here in Canada. You mean "propagation" from the verb to propagate or to reproduce something. Also not "and" existing one but "an".

    And the correct word for your title is "existing" not "existant" (which in any case should be spelled existent).

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  2. 2:00am drafting and proofing. Likely more. Tks much...

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  3. Excellent examination of the Ginga Rubra genotype, Alan. I found it interesting that, shortly after adding your Shimmelpfennig Platinum crossover gups to my collection, I also added a couple of Ginga Rubra gups from another source. I did so based on the phenotype BOTH strains exhibit to the eye. You have confirmed that intuitive inkling I had; they are the same or similar phentoypes, indeed. Of course, your knowledge stems from the use of the scientific method. Mine stems from years of being a "sight" breeder of many species. Sometimes I am correct in my assumptions BEFORE actually making crosses and examining the results. In other instances I am not even close. My grandfather taught me that while teaching me how to hybridize irises and tulips. He noted that a good breeder, whether of plant or animal, must have a good imagination. The ability to imagine what a cross might produce is, in my estimation, one of the hallmarks of a good "sight" breeder. Your examination of the Ginga Rubra phenotype, however, will hasten my success in the crosses I have planned with Endler's LBs for the Shim Plats and the Ginga Rubras.

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