Once in a Blue Moon

One of the best-known facts of genetics is that a cross between a horse and a donkey produces a mule. Actually, it's a cross between a female horse and a male donkey that produces the mule; the reciprocal cross, between a male horse and a female donkey, produces a hinny, which has smaller ears and a bushy tail, like a horse ( FIGURE 9.1). Both mules and hinnies are sterile because horses and donkeys are different species with different numbers of chromosomes: a horse has 64 chromosomes, whereas a donkey has only 62. There are also considerable differences in the sizes and shapes of the chromosomes that horses and donkeys have in common. A mule inherits 32 chromosomes from its horse mother and 31 chromosomes from its donkey father, giving the mule a chromosome number of 63. The maternal and paternal chromosomes of a mule are not homologous, and so they do not pair and separate properly in meiosis; consequently, a mule's gametes are abnormal and the animal is sterile.

In spite of the conventional wisdom that mules are sterile, reports of female mules with foals have surfaced over the years, although many of them can be attributed to mistaken identification. In several instances, a chromosome check of the alleged fertile mule has demonstrated that she is actually a donkey. In other instances, analyses of genetic markers in both mule and foal demonstrated that the foal was not the offspring of the mule; female mules are capable of lactation and sometimes they adopt the foal of a nearby horse or donkey.

In the summer of 1985, a female mule named Krause, who was pastured with a male donkey, was observed with a newborn foal. There were no other female horses or donkeys in the pasture; so it seemed unlikely that the mule had adopted the foal. Blood samples were collected from Krause, her horse and donkey parents, and her male foal, which was appropriately named Blue Moon. A team of geneticists led by Oliver Ryder of the San Diego Zoo examined their chromosomal makeup and analyzed 17 genetic markers from the blood samples.

I 9.1 A cross between a female horse and a male donkey produces a mule; a cross between a male horse and a female donkey produces a hinny.

(Clockwise from top left, Bonnie Rauch/Photo Researchers; R.J. Erwin/Photo Researchers; Bruce Gaylord/Visuals Unlimited; Bill Kamin/Visuals Unlimited).

Krause's karyotype revealed that she was indeed a mule, with 63 chromosomes and blood type genes that were a mixture of those found in donkeys and horses. Blue Moon also had 63 chromosomes and, like his mother, he possessed both donkey and horse genes ( FIGURE 9.2). Remarkably, he seemed to have inherited the entire set of horse chromosomes that were present in his mother. A mule's horse and donkey chromosomes would be expected to segregate randomly when the mule produces its own gametes; so Blue Moon should have inherited a mixture of horse and donkey chromosomes from his mother. The genetic markers that Ryder and his colleagues studied suggested that random segregation had not occurred. Krause and Blue Moon were therefore not only mother and son, but also sister and brother because they have the same father and they inherited the same maternal genes. The mechanism that allowed Krause to pass only horse chromosomes to

Donkey 2n = 62

-rO Horse

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