Testing Adaptive Models Of Sex

Allocation in a Polymorphic Species

(c) 2003 Doug Von Gausig

Introduction
The first theories concerning evolution and sex ratio were introduced by Darwin (1871) in his ground-
breaking book The Descent of Man and Selection in Relation to Sex. One of his most prevalent theories
is that a facultative, or changing, sex ratio is a direct product of natural selection.
Methods
Sample Collection
Approximately 50% of the world’s species of birds, including the white-throated sparrow, are not
sexually dimorphic. Blood samples were collected for sexing through molecular DNA analysis.
- All blood sample collection was done by Tuttle and field teams.

Two major classes of adaptive hypotheses were tested in this study:

Fisher’s Hypothesis
* Homeostasis - sex ratio should be maintained at ideal equlibrium ratio of 0.5
- any biased sex ratio at birth will be in favor of the rarer sex
* Cranberry Lake Biological Station, Aidirondack Mountains, NY
* Indentified by a serial number and band colors
* This study encompasses data collected in 2000-2004.
* 282 chicks were analyzed for a composite total of 92 clutches.

Trivers-Willard Hypothesis Molecular Analysis

* Better conditioned parents should invest in the sex that has the best chance of
A DNA-based sex identification method was used utilizing two CHD (chromo-helicase-DNA-
producing the most grand-offspring.
binding) genes, CHD-W and CHD-Z, located on the avian sex chromosomes W and Z, respectively.

The White-Throated Sparrow 1) Blood was separated through centrifugation and red blood cells (RBCs) were stored.
(Zonotrichia albicollis)
2) DNA was extracted using the Promega DNA IQ System.
The white-throated sparrow , is a passerine which inhabits northeastern United States and Canada.
3) P8 and P2 primers were used to amplify the needed sequence through PCR.
They are a unique avian species in that there is a separate genetic polymorphism that is connected with
4) PCR product loaded into the ABI Prism 310 Genetic Analyzer, capillary electrophoresis.
morphological and behavioral distinctions. These are caused by a pericentric inversion on the second
chromosome, 2m. A physical distinction occurs in plumages; without the presence of the inversion
(homozygous 2/2), the bird has a tan stripe on its head, but with the inversion (heterozygous 2/2m),
the stripe is white.

White-throated sparrows mate

disassortatively, meaning, naturally,
tan females mate with white males
while white females mate with tan
males.

Tan White Statistical Analysis

Parental Promiscuous *Calculated proportion of males in each clutch
The inversion causes variation in behavior where tan morphs tend to take up a more parental role in *Arcsin square root transformation
caring for offspring while whites are much more socially aggressive and sexually promiscuous. White *ANOVA
males often seek extra-pair copulations (EPCs) with neighboring females that are not their social mates. *Year, pair type, hatch date
In an earlier study tan males were found not only to be more attractive than white males, but also *Post hoc Tukey-Kramer HSD, ����
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=0.05
tan males and their white female mates (TxW) were found to nurture their offspring and contribute * JMP 4.0.4 statistical analysis program
parental investment much more than white male/tan female parents (WxT).
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The objective of this study was to test the Fisher and the Trivers-Willard hypotheses in white-throated
sparrows.
Predictions
Results
Fisher’s Hypothesis: * Sex ratio of WxT clutches do not vary significantly
Clutch sex ratios will be facultative in order to maintain the ideally stable from year to year (F4,36=0.67, p=0.62)
proportion of 0.5. Regardless of parent pair type, trends will be the same * Sex ratio of TxW clutches vary significantly
through out. from year to year (F4,49=3.61, p=0.01)
* Overall average sex ratio of the population did
Trivers-Willard Hypothesis: not differ significantly from 0.5.
Clutch sex ratios will differ between TxW parents and WxT parents. Trends
should correlated to difference in pair type based upon parental effort.

Abstract
* TxW clutch ratios do not differ significantly in early
and late clutches (F1,52=1.30, p=0.25)
* WxT clutches produce a significantly lower
proportion of males in late clutches
(F1,38=4.96, p=0.03)

Discussion Fisher’s Hypothesis:

* Overall, not significantly different from 0.5
* Clutch sex ratios of TxW≠WxT

Trivers-Willard Hypothesis:
* The trends in clutch sex ratio of TxW pairs
and WxT pairs differ.
White male/tan female parents showed evidence that they are able to facultatively shift the sex
ratio of their clutch in response to environmental cues such as predators. The later breeding season
harbors many more predators and a greater risk of depredation in nests. Because TxW pairs are better
parents, they need not shift their sex ratio; they are able to fend off predators. On the other hand,
with more predators, WxT pairs have a disadvantage as they are “poorer” parents. Therefore, in order
to increase their chances of having some grand-offspring, they produce more daughters, instead of
more expensive sons. This strongly supports the Trivers-Willard hypothesis. In additional support, the
data also show that TxW parents facultatively shift their sex ratio from year to year.
References
* Darwin, C. 1871. The Descent of Man and Selection in Relation to Sex. London: John Murray. Future Directions
* Ellegren, H. 1996. First gene on the avian W chromosome (CHD) provides a tag for universal In the future I hope to look further into parental investments in nestling sex ratio, since the parental
sexing of non-ratite birds. Proceedings of the Royal Society of London B 263: 1635-1641. condition and consequent amount of investment is very distinguished throughout the data. Another
* Fisher, R. A. 1930. The Genetical Theory of Natural Selection. Oxford: Clarendon. interesting aspect to look at would be the effects of laying order on the sex of the nestling. In addition,
* Trivers R. L. and Willard D. E. 1973. Natural selection of the parental ability to vary the sex I hope to continue this study in order to further examine cyclic patterns.
ratio of offspring. Science 179: 90-92.

*All photos of live birds were taken by the Tuttle field team.
*All others, unless otherwise credited are original photos.