It is obvious that many birds exhibit sexual dimorphism of color. But the causal
mechanisms resulting in sexual dichromatism have been well demonstrated for only a
few species such as chickens and finches. Sex hormones control the plumage color and
form in the brown leghorn (Greenwood and Blyth, 1938). Witschi (1935, 1937) and
students have shown that within the weaver finches, three kinds of control are involved:
sex hormones, gonadotrophic hormones, and genetic constitution.
The auto-sexing varieties of chickens and geese (Hutt, 1949; Quinn, 1939) and of
domestic pigeons reveal sex dichromatism by the time of hatching and in the first plum-
age. Auto-sexing refers to a genetic method of detecting sex in birds and requires a sex-
linked gene with dosage effects easily detected early in the birds' development. A wild
species exhibiting auto-sexing is the Snow Bunting, Plectrophenax nivalis (Pitelka,
personal communication). Auto-sexing is difficult to explain by known hormonal mecha-
nisms, and relatively direct gene control of such sexual dichromatism is indicated
(Hollander, 1942);
Among the approximately 14 species of doves in the genus Streptopelia only one, the
Dwarf Turtle Dove (Streptopelia tranquebarica), exhibits marked sexual dichromatism.
This dove is native to the Philippines, southeastern Asia, and India. Immature and
female tranquebarica are "gray." The mature male has a bluish head, with the color
extending to the neck ring, and a reddish brown "back."
Several auto-sexing pigeons and tranquebarica were available from a pigeon colony
housed in a barn by the Department of Genetics at the University of Wisconsin, Madi-
son, Wisconsin, and since they seemed to represent opposite extremes of early and late
dichromatism within the Columbidae the following experiments with hormones were
conducted.
The data on the domestic pigeon or Rock Dove (Columba livia) were collected in
connection with another study (Miller and Wagner, 1955) in which the hormones were
donated by Dr. W. H. McShan and the implants and some of the injections were made
by Frederic H. Wagner.
MATERIALS AND ETItODS
The sex of the mature Dwarf Turtle Doves was ascertained by their color. This
diagnosis agreed with the method of sexing described by Miller and Wagner (1955)
and Hanson (1953). Female doves were injected with testosterone propionate, the males
with estradiol benzoate, and the controls were injected with sesame oil (the diluent for
the hormones) according to the following plan:
Dove Sex
D801.136 Male
D801.134 Female
D801.112 Male
DS01.140 Female
D801.171 Female
Daily injection intramuscularly
0.1 ml. sesame oil
0.1 ml. sesame oil
0.1 ml. estradiol benzoate (0.166 rag.)
0.02 ml. testosterone propionate (0.1 rag.)
0.08 ml. testosterone propionate (0.4 rag.)
On the third day after injections were begun, some feathers were plucked from the
right half of the bird's head and from the right wing. The wing feathers plucked were
lesser secondary and marginal coverts. Eleven days after plucking, the last injections
were given to the controls and to D801.171 since the regenerating feathers were already
showing their color. D801.112 and D801.140 received their last injections 20 days
and 28 days after plucking, respectively, since their feather regeneration was slower.
D801.171 died 18 days after plucking, but most of the regenerating feathers were about
two-thirds regrown and were saved. The head feathers of 1)801.112 were extremely de-
layed in regeneration and very probably had no effect of the injected hormone since
the injections had been stopped at least 30 days previously.
The feathers grown before and after treatment were compared and colors checked
with Ridgway's "Color Standards and Color Nomenclature."
Auto-sexing strains of the domestic pigeon have been made utilizing the mutant
"faded," St r, which has been described by Hollander (1942). The faded gene is sex-
linked and causes sparse down and absence of pigment on the bills of homozygous males
just hatched but permits nearly normal down and pigment on the bills of hemizygous
females. The first feathers of the homozygous faded male squabs are white with rare
flecks of color, usually black but often reddish in color. The first plumage of a homozy-
gous faded male does not differ in color from later plumages except that the flecks may
become more numerous and larger. The hemizygous faded female has nearly normal-
colored plumage but has a "washed out" appearance.
Five-week-old, opposite-sexed siblings from a pure line of faded pigeons received
intramuscular injections daily of the opposite sex hormones0.25 ml. of testosterone
propionate was given the female (equivalent to 12.5 mg. per injection and equal on a
weight basis to 0.04 ml. or 0.2 mg. if given to a Dwarf Turtle Dove), and 0.5 mi. of
estradiol benzoate (0.83 mg.) was given the male.
Some tail feathers, secondaries from the right wing, and feathers from the back were
plucked from each bird at the first injection of hormone, and these of course were saved
for comparison with the regenerated feathers. Injections continued for 18 days until the
regenerating feathers were nearly fully regrown.
Two additional males six weeks of age were implanted with diethylstilbesterol pel-
lets; they were similarly plucked and the feathers compared.
RESULTS
Several birds other than those listed were given different doses of the hormones at
the beginning of the experiment. These birds died in a Newcastle epidemic before results
could be obtained (see Hanson and Sinha, 1952, and Miller and Wagner, 1955).
The faded female of Columba livia injected with testosterone propionate developed
typical male doacal papillae during the first week of injections, that is, at six weeks of
age (Miller and Wagner, 1955). Papillae are not ordinarily distinguishable until males
are four to six months old. No color change was observed in any of the regenerated
feathers of the injected or pellet-implanted pigeons when these were compared to the
original plucked feathers. There was a slight increase of black flecks in the males' feath-
ers but this is normal for such males.
All Dwarf Turtle Doves receiving the "reverse" sex hormone showed intermediate
color and pattern of feathers tending to be much more like the opposite sex. As Salo-
monsen (1940) has indicated, the influence of hormones on feather color and pattern is
not "all or none" but often may be intermediate. New feathers of the control birds
1)801.134 and 1)801.136 were one grade darker than the previous old feathers, except
for the male wing color which showed little difference. This kind of change had been
noticed for new feathers of this and other species of doves, probably resulting from
sun fading or weathering of the older feathers.
Comparisons of the feather color of the Dwarf Turtle Doves are given in table 1.
Specimen number and sex
D801.134 9 control
D801.136 control
D801.112
0.1 cc. estradiol
D801.140 9
0.02 cc. testosterone
D801.171 9
0.08 cc. testosterone
Table 1
Results of Reversing Sex Hormone on Streptopelia tranquebarica
Standardization of untreated feathers Standardization of regrown feathers
Wing: army brown edges deepening natal brown (next grade to army
gradually to deep mouse gray in brown) deepening to deep
the proximal half of the feather mouse gray
Head: deep mouse gray dark mouse gray lightening to
deep mouse gray
Wing: Hay's brown changing abrupt- Hay's brown changing abruptly
to deep mouse gray (no change)
ly to deep mouse gray in the proxi-
mal half of feather
Head: dark gull gray
Wing:
Head:
Wing:
than in D801.134
Head: color same as in D801.134
Wing: colors same as in 1)801.134
slate gray (next darker grade to
dark gull gray)
colors same as in D801.136 Hay's brown edges* only, grading
to deep mouse gray
colors same as in D801.136 (delayed regeneration of feathers)
no change, dark gull gray
army brown edges but larger very large area of the edges Hay's
brown with an abrupt change to
deep mouse gray in the proximal
part of the feather
dark gull gray
feathers "two-thirds" regrown;
l-lay's brown slightly "peppered"
with and deepening to deep
mouse gray
Head: colors same as in D801.134 no feathers; bird died
* Italics indicate hormonal influence.
Standardization was made twice under fluorescent and "ordinary" bulb lighting using Ridgway's "Color Standards
and Color Nomenclature."
DISCUSSION
In spite of the loss of some birds before results could be obtained, it has been demon-
strated that sexual dichromatism in Streptopelia tranquebarica is under the immediate
control of the sex hormones. In the auto-sexing Columba livia, "direct" gene action
without sex hormonal influence is indicated as the cause of sexual dichromatism.
It has been shown that sex hormones control the plumage form and color in brown
leghorns (see Juhn, Faulkner and Gustavson, 1931; Domre, Gustavson and Juhn, 1932;
and Greenwood and Blyth, 1938). Sex hormones alone do not explain sexual dichro-
matism in all birds even for those which show marked hormone influence (Danforth,
1937). Half-and-half or sectorial mosaics, male on one side and female on the other,
with regard to color are known (Hollander, 1944), and species showing seasonal plumage
changes complicate the problem. Witschi (1935) and students have explained some of
the complications in their work on weaver finches, in which a complex interaction of sex
hormone, gonadotrophic hormone, and genetic constitution is shown. Also see Miller
(1935) for some effects of thyroxin.
Much work remains to be done to clarify the mechanisms of sexual dichromatism.
For example, what is the genetic basis for the differences of S. tranqebarica from other
species of Streptopelia that enable it to exhibit sex hormonal control of its sexual di-
chromatism? Certainly the genetic interactions with the environment both internal
(especially hormones) and external can be complex.
SUMMARY
Sexual dichromatism in two species of the Columbidae exemplify the two extreme
types of control: "direct" genetic control, and sex hormonal control.
Reversal of the predominant sex hormone by injection of testosterone propionate or
estradiol benzoate in mature individuals of the Dwarf Turtle Dove (Streptopelia tran-
quebarica) resulted in a corresponding change in the feather color in the head and wing
which in this species shows sexual dichromatism. Control tranquebarica, male and fe-
male, injected with sesame oil, showed no change.
Reversal of the predominant sex hormone in the auto-sexing domestic pigeon
(Columba livia) had no effect on the plumage color.
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