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4.2 Sex Chromosomes and Sex Determination    97


                           TABLE 4.1      Sex Determination in Fruit Flies and Humans

                                                           Complement of Sex Chromosomes
                                     XXX           XX            XXY             XO            XY            XYY        OY

                        Drosophila   Dies      Normal female   Normal female   Sterile male   Normal male   Normal male   Dies
                        Humans       Nearly    Normal female   Klinefelter   Turner female   Normal male   Normal or    Dies
                                     normal                   male (sterile);   (sterile); short,         nearly normal
                                     female                   tall, thin     webbed neck                  male
                       Humans can tolerate extra X chromosomes (e.g., XXX) better than can Drosophila because in humans all but one X chromosome becomes a Barr body, as discussed
                       later in this chapter. Complete absence of an X chromosome is lethal to both fruit flies and humans. Additional Y chromosomes have little effect in either species.
                       Although the Y chromosome in Drosophila does not determine whether a fly looks like a male, it is necessary for male fertility; XO flies are thus sterile males.

                       Species Vary Enormously in Sex                           TABLE 4.2      Mechanisms of Sex
                       Determining Mechanisms                                                  Determination
                       Other species show variations on this XX versus XY chro-                     ♀                ♂
                       mosomal strategy of sex determination. In fruit flies, for
                       example, although normal females are XX and normal    Humans and             XX               XY
                                                                             Drosophila
                       males XY (see Fig. 4.2), it is ultimately the number of X
                                                                                                    XX
                       chromosomes (and not the presence or absence of the Y)   Moths and      (hermaphrodites       X0
                       that determines sex. The different responses of humans and   C. elegans   in C. elegans)
                       Drosophila to the same unusual complements of sex chro-
                       mosomes (Table 4.1) reveal that the mechanisms for sex   Birds and          ZW                ZZ
                                                                             Butterflies
                       determination differ in flies and humans. XXY flies are
                       female because they have two X chromosomes, but XXY   Bees and             Diploid          Haploid
                                                                             Wasps
                       humans are male because they have a Y. Conversely, be-
                       cause they have one X chromosome, XO flies are male,   Lizards and          Cool             Warm
                       while XO humans are female because they lack a Y.     Alligators         temperature       temperature
                          The XX = female / XY = male strategy of sex determi-  Tortoises and      Warm             Cool
                       nation is by no means universal. In some species of moths,   Turtles     temperature       temperature
                       for example, the females are XX, but the males are XO. In C.   Anemone Fish   Older adults   Young adults
                       elegans (one species of nematode), males are similarly XO,   In the species in the top three rows, sex is determined by sex chromosomes. The
                       but XX individuals are not females; they are instead self-   species in the bottom four rows have identical chromosomes in the two sexes,
                       fertilizing hermaphrodites that produce both eggs and sperm.   and sex is determined instead by environmental or other factors. Anemone fish
                                                                           (bottom row) undergo a sex change from male to female as they age.
                       In birds and butterflies, males have the matching sex chro-
                       mosomes, while females have an unmatched set; in such spe-
                       cies, geneticists represent the sex chromosomes as ZZ in the   and probably many other traits as well. Sutton and other
                       male and ZW in the female. The sex having two different sex   early adherents of the chromosome theory realized that the
                       chromosomes is termed the heterogametic sex because it   perpetuation of life itself therefore depends on the proper
                       gives rise to two different types of gametes; conversely, the   distribution of chromosomes during cell  division. In the
                       sex with two similar sex chromosomes is the homogametic   next sections, you will see that the behavior of chromo-
                       sex. The gametes of the heterogametic sex would contain   somes during mitosis and meiosis is exactly that expected
                       either X or Y in the case of male humans, and either Z or W   of cellular structures carrying genes.
                       in the case of female birds; the gametes of the homogametic
                       sex would contain only an X (humans) or only a Z (birds).   essential concepts
                       Yet other variations include the complicated sex-determination
                       mechanisms of bees and wasps, in which females are diploid   •  Many sexually reproducing organisms have sex
                       and males haploid, and the systems of certain fish, in which   chromosomes that are sex-specific and that determine sex.
                       sex is determined by changes in the environment, such as   •  In humans, male sex determination is triggered by a
                       fluctuations in temperature. Table 4.2 summarizes some of   Y-linked gene called SRY; female sex determination
                       the astonishing variety in the ways that different species have   occurs in XX embryos by default.
                       solved the problem of assigning sex to individuals.   •  Mechanisms of sex determination vary remarkably; in
                          In spite of these many differences between species,   some species sex is determined by environmental factors
                       early researchers concluded that chromosomes can carry   rather than by specific chromosomes.
                       the genetic information specifying sexual characteristics—
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