[Front matter only] The Guppy Color Manual: Explorations of Guppy Color Biology and Genetics (aka The Guppy Color Manual: Explorations in Guppy Color Biology and Genetics)

14 Pages·2016·0.38 MB·English

by  Philip Shaddock

The Guppy
Color Manual

Explorations of Guppy Color Biology and Genetics

First Edition
June 2012
The Guppy Color Manual: Explorations in Guppy Color Biology and Genetics
© 2012 Philip Shaddock
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10 9 8 7 6 5 4 3 2 1
Contents
Foreword 9
Preface 11
Acknowledgements.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .13
1 The Study of Guppy Color 15
The.Birth.of.Guppy.Genetics.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .15
Mining.the.Data ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... .15
The.First.Guppy.Color.Cell.Paper... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 16
Developmental.Biology.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .18
The.Discovery.of.Gene.Regulation ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... .18
Guppy.Color.Layering.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .19
The.Fourth.Dimension. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .22
The.Multipotent.Color.Cell.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .22
Puppets.on.A.Stage.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .23
Evo-Devo:.A.New.Synthesis... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 24
2 The Color Cells 27
Color.and.Water.Depth. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .27
The.Skin... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 30
The.Color.Cells... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 32
Two.Different.Embryonic.Origins.of.Color.Cells... .. .. .. .. .. .. .. .. .. .. .. .. .. .. 33
Color.Cell.Types. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .36
Color.Cell.Biochemistry. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .36
Pigment.Color.Cells.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .38
Melanophores.(Black.Color.Cells). .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .38
Dermal.Melanophores. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .41
Types.of.Melanophores. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .42
Melanosomes. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .44
Pigment.Color.Cells:.Red.and.Yellow. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .44
Carotenoids.and.Pteridines. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .46
Guppy.Orange. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .47
Pteridine.Biosynthesis. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .47
Pigment.Blues.Versus.Reflective.Blues. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .48
Guppy.Sun.Tan.Lotion.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .49
The.Structural.Color.Cells.of.the.Guppy ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... .49
Common.Types.of.Iridophores ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... .52
Comparing.Leucophores.and.Iridophores. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .53
Reflecting.Platelets.of.the.Iridophores. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .56
Mosaic.Color.Cells. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .58
Color.Cell.Motility.(Chameleon.Effect). .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .59
Hormonal.and.Nervous.Control.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .60
Chameleon.Color... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 60
Hormonal.Factors.in.Color.Change ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... .61
Color.Change.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .61
Color.Cells.Result.From.Biological..Pathways. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .63
Color.Cell.Development.and.Differentiation... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 65
Other.Factors. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .66
Pigmentation.Anomalies ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... .67
3 Color Cell Layering 69
The.Bagnara.Dermal.Chromatophore.Unit. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .70
Filtering.Layer. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .71
Reflecting.Layer.... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 72
Absorbing.Layer ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... .73
Altering.the.Saturation.or.Intensity.of.a.Color. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .74
Color.Cells.Interact.to.Produce.Novel.Color. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .74
Pastel.Colors.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .75
Structural.Colors... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 76
Blue.and.Silver.Iridophores. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .76
Blue.Iridophores ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... .79
The.Interplay.Between.Iridophores.and.Red.Color. .. .. .. .. .. .. .. .. .. .. .. .. .. .83
Tyndall.Blue... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 84
Melanistic.Guppies... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 85
4 The Wild-Type Guppy Patterns 89
Guppy.Reticulation.Pattern. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .89
Melanophores.and.the.Reticulation.Pattern.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .91
The.Reticulation.Pattern.and.the.Fins.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .96
Male.Polymorphic.Color.Pattern... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 99
The..Wingean.Pattern.is.Strictly.Y-Linked. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..100
Winge.treated.the.patterns.as.two.dimensional.patterns.(Figure.18) . ... ... ... ..100
Winge.and.XX.Males. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..103
The.Wingean.Supergene. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..104
Wild-Type.Patterns.Highly.Homozygous.Across.Loci.... .. .. .. .. .. .. .. .. .. .. .. .. .108
Summary... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .109
5 Pattern Theory 111
Bagnara’s.Review.of.Pattern.Theory. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..111
Color.Pattern.Formation.in.Zebrafish. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..112
Hormonal.Influences.on.the.Development.of.Pigmentation.Patterns.. .. .. .. .113
Fujii.on.Patterns ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ..113
Kelsh.on.Patterns. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..114
Other.Factors.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .115
What.Other.Animal.Studies.Teach.Us.About.Patterns.. .. .. .. .. .. .. .. .. .. .. .. .. .115
Fundamental.Design... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .116
Variations.in.Number.and.Kind. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..117
Symmetry.and.Polarity... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .118
Spemann.Organizer. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..118
Master.Building.Genes:.Hox.Genes... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .120
The.Guppy.Segmented.Body. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..123
The.Segmented.Body:.Fin.and.Color.Patterns. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..128
Mapping.the.Body... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .128
The.Geography.of.the.Body.(and.Patterns!)... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .128
Lateral.Inhibitions. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..132
Patterns.and.Genetic.Switches.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .133
Combinatorial.Logic.of.Switches... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .138
Patterns.Are.the.Whole.of.a.Sum.of.Parts... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .140
Conclusion. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..142
6 Guppy Black: Hypomelanism 145
The.Story.So.Far . .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..145
Guppy.Polymorphism.and.Black... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .147
Hypomelanism. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..150
Melanosynthesis.Fault:.Albinism... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .150
WREA.Guppies... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .152
Melanophore.Migration.Faults.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .153
The.Blond.and.Golden.Mutations. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..158
The.Golden.Mutation... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .158
The.Blond.Mutation. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..163
Bleach.Blonds. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..165
The.NII.Gene... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .165
The.Cream.Double.Recessive.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .167
7 Guppy Black: Melanism 169
Moscow.Blue.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .175
Moscows.Are.a.Simple.Combination.of.Wild-Type.and.the.Moscow.Gene(s)... .176
The.Onyx.Gene.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .178
The.Midnight.Gene.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .180
NiI.and.NiII.in.the.Scientific.Literature.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .184
The.“Half.Half-Black”.Pattern.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .188
A.Biological.Basis.for.Color.Suppression.in.the.Guppy.Anterior?... .. .. .. .. .. .. .. .190
Is.there.a.genetic.type-switch.between.melanophores.and.leucophores?. .. .. .. ..193
8 Red and Yellow Mutants 201
The.Full.Red.Debate. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..204
Full.Red.Crossing.Experiments.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .205
F1.Full.Red.Male.to.Wild-Type.Female.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .206
The.Glass.belly.Gene.and.Full.Red ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ..208
The.Blaus ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ..210
European.blau... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .210
European.versus.Asian.Blau. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..211
Asian.blau... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .212
The.Effect.of.the.Blau.Mutations.on.Guppy.Patterns. .. .. .. .. .. .. .. .. .. .. .. ..214
Blau.Removes.Red,.Does.Not.Substitute.Blue. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..214
Black.and.Blau.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .215
Dosage.Effects... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .218
Asian.blau.Fin.Development... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .219
Asian.blau.As.a.Color.Tool.in.Guppy.Genetics. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..219
Heterozygous.Blau.Moscow. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..219
Asian.blau.Genetic.Notation... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .222
The.Magenta.Mutation... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .222
Magenta.Description ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ..222
The.Magenta.genetic.Switch.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .224
Fin.Shape... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .227
Magenta.Notation ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ..227
9 The Striped Guppy 229
Snakeskin.Cells.Under.the.Microscope. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..231
Snakeskin.as.“Disorganized”.Reticulation.Pattern. .. .. .. .. .. .. .. .. .. .. .. .. ..233
Colors.and.the.Snakeskin.Pattern.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .236
Snakeskin.Caudal.fin.Patterns ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ..239
Research.Information.on.Patterns. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..241
Lace.Genetics. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..245
Half-Black.Snake.Hybrids ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ..246
Dorsal.Length.in.Snakeskins... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .246
Peduncle.Vertical.Bar.Pattern. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..247
The.Zebrinus.Gene. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..250
The.Tigrinus.Gene. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..251
Is.There.a.Bar,.Zebrinus.or.Tigrinus.Gene?. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..252
Other.Spots.in.Snakeskin.Patterns ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ..252
Grass.Guppy... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .253
Grass.Genetics ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ..257
Snakeskin.Breeders.Comments.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .257
10 Fin Shape and Color 259
Wild.Type... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .260
Fin.Anatomy.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .262
Wild.Type.Fin.Shapes... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .264
Caudal.Fin... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .264
Dorsal.Fin. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..265
Pectoral.Fins... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .265
Pelvic.Fins... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .265
Anal.Fin... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .265
Basic.Fin.Genetics. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..265
Speculating.About.Fin.Shape.Factors... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .267
Testosterone.and.Fin.Extensions... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .269
Fin.Extensions.and.Color. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..270
The.Fin.Shapes.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .270
The.Swords. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..270
Sex-Reversed.Swordtail.Female. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..275
Pintail.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .276
Elongated.Dorsal.(Hi-Fin) ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ... ..276
Fantail.Fin.Shape... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .277
Merah.Fin.Shape... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .277
Ribbon.or.Giessen.Fin.Shape.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..278
Roundtail.Fin.Shape. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..280
Spadetail.(Coffertail).Fin.Shape. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..280
Speartail.Fin.Shape... .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .281
Swallow.(Berlin).Fin.Shape. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..283
Breeding.the.Swallows. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..284
A References 287
B Guppy Dictionary 291
Other.Guppy.Designer.Books.
. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..301
| 9
Foreword
Philip Shaddock decided to systematically document how guppy color genes interact with each other.
This fundamentally important topic was not adequately treated by any scientific text or hobbyist
reference book. To be sure, many individual interactions had been reported, but no systematic study
could be found. Years later, the results of these studies, often done and analyzed by the author himself
with frequently added information, results and interpretations gleaned from others, is now in print.
This monograph is an authoritative textbook on guppy color biology. It assumes a knowledge of basic
guppy genetics, which is very well covered in the author’s earlier books “Guppy Genetics Simplified”
and “The Theory and Practice of Guppy Breeding”. This book develops the more complex concepts
needed to really understand the biology of guppy pigmentation. A vast range of different color genes
and their interactions is described and explained. This book is a handy reference to both the guppy
hobbyist and professional guppy breeder. In fact, I believe that no serious guppy hobbyist or breeder
can afford to miss this book! It is also the only reference I know of that will be equally valuable to the
scientist who is interested in guppy pigment biology. Every researcher interested in fish pigment sys-
tems will want a copy of this book as well.
The author refers to earlier work showing that many Y-linked genes are maintained together within
“Wingean super genes” on the Y chromosome. This led him to construct his own significant hypothesis
regarding guppy color genetics. He has suggested that most of the color genes that code for structural
proteins are located on the autosomes (non-sex chromosomes) rather than on the sex chromosomes
themselves. His unique suggestion is that the sex chromosomes (and the Y-chromosome in particular)
contain genes that regulate these actual color genes that are found on the autosomes. This is a sig-
nificant shift in the scientific concept of which genes are sex-linked as opposed to being autosomally
linked. I am confident that this hypothesis will be investigated by molecular biologists in the years to
come.
This book contains a wealth of information for the reader regardless of their level of expertise. Basic
concepts and hard-to-find journal articles are described so that any one can understand them. Philip’s
training as an English major makes itself evident. This well written book will be enjoyed by hobbyist
and scientist alike. I learned a lot reading the manuscripts, and enjoyed every minute!
Richard Squire, Ph.D. (Genetics)
Retired Full Professor of Biology
University of Puerto Rico – Mayaguez Campus
10 | Foreword