Tuesday, February 17, 2015

Sex determination in birds

Humans, other mammals, and some insects genetically determine sex via X and Y chromosomes. In the XY sex-determination system, females inherit an X chromosome from each parent, while males inherit an X-chromosome from their mothers and a Y-chromosome from their fathers. Since homogametic XX females produce only X chromosomes, heterogametic XY males dictate the sex of their offspring.

Human watchers (including myself) often apply sex-based characteristics to our beloved eagles. Mom and Dad appear to have many of the same housekeeping arrangements and arguments that we do, so it might be surprising to learn that they turn our XY sex-determination system on its head!

Introducing...the ZW sex-determination system!
Birds, some reptiles, some amphibians, and some fish replace our X and Y with the ZW system. Let's look at a few differences:
  • I'm a female homogametic XX and my husband is a male heterogametic XY, but female birds are heterogametic ZW and male birds are homogametic ZZ. Among other things, this means that Mom's ova determines the sex of the little E's (or D's) we coo over in Decorah and elsewhere. 
  • XY and ZW chromosomes come from different areas of the genome. That means that our system for genetically determining sex is not very closely related to a ZW'er's system for determining sex, even though they both work roughly the same way. 
  • ZW sex determination is dose-dependent, at least in birds. Z doesn't create a male in and of itself. Two Z's (ZZ) are required to make enough male-determining product. Females may be female not because of the W, but because they lack two Z's.
Having said that, ZW females still produce eggs and ZZ males still produce sperm. There are some other similarities between us and them:
  • The non-recombining Y and W chromosomes have both degenerated over time. Today, the mammalian X carries over three times more genes than the Y does, whereas the chicken Z carries over ten times more than the W. Like our Y-chromosome, the W doesn't combine well with its opposite. Deletion, degradation, and mutations occur more quickly on the Y and W than on the X and Z. 
  • Y and W resemble X and Z more closely in 'primitive' animals and plants than in those that have evolved more recently, and both Y and W continue to shrink. Will human maleness and bird femaleness disappear? I don't think it's likely, but some people believe that new forms of sex determination might arise from somewhere else in the genome.  
How does the ZW system affect birds?
We know birds aren't human, even though we might call them Mom and Dad, think of them as friends, and invest them with human-like feelings, duties, and housekeeping systems. Still, it seems odd that they have a system of genetically-determined sex that is so different than ours. How does the ZW system affect them? 
  • Z chromosomes contain more genetic information than W chromosomes. Since ZZ male birds have two Z chromosomes, they are more likely to pass on sex-linked traits than ZW female birds. For example, Decorah daughter (ZW) Four got a Z from Dad and a W from Mom, while son (ZZ) Decorah got a Z from each parent. A male bird contributes a Z to sons and daughters, while a female passes a Z to sons only. Since Dad's Z contains a lot more information than Mom's W, Dad contributes more genetic information on that particular chromosome, especially when it comes to daughters. In humans, females conserve more sex-linked traits than males, and pass them on to male and female children. The opposite is true in birds. 
  • What about that flashy male plumage? We know that male birds conserve more sex-linked traits and pass them on to sons and daughters, but male plumage is more complicated than it appears.  Recent work published in the journal Evolution indicates that female birds were once as flashy as males. We think that sexual selection drove male color evolution (females prefer colorful males), and natural selection drove female loss of color (brighter females and young were more likely to be spotted by predators and competitors). 
  • So what role did the ZW system play? A couple of ideas. W has degraded over millions of years and carries much less information than it did previously. If the gene for female color was located on W, it may have been lost somewhere in the past. Alternatively, Z is more highly expressed than W. The resulting imbalance in hormonal secretions between lower ZW females and higher ZZ males drives fundamental sex differences in phenotype, development, and physiology. A ZW chromosome may not provide enough oomph to turn bright plumage on elsewhere in the genome, assuming it exists in female birds.
  • What about birds of prey? Unlike most birds, female raptors or birds of prey are larger than male birds of prey. And unlike most birds, male and female raptors sport the same plumage. With the exception of kestrels, we can't use plumage color to tell males and females apart. What happened to de-link plumage color with sex and why is size, which is still linked to sex, inverse almost uniquely among birds of prey? Is it expressed in the Z or W chromosomes (W chromosomes are 'large in many raptors'), or elsewhere in the genome? Curious minds want to know!  
Why do we have XY and ZW?
No one knows for sure. All snakes and all birds use ZW determination, and all mammals use XY determination (even when it gets weird, as it does with platypus and voles, X is always present). But fish, amphibians, turtles, and lizards might use XY, ZW, or temperature-dependent selection (TDS) depending on the species. The switch from a variety of systems (ZW, TDS, XY) appears to have occurred around the time the ancestor of all reptiles split from the ancestor of all mammals, but it's not yet clear whether there was an intermediate stage between ZW and XY systems, or whether the transitions occurred directly.  

Is the ZW system really that different? I don't think so. While lovely heterogametic ZW Mom determines sex and handsome homogametic ZZ Dad passes more genetic material to his offspring, males still produce sperm, females still produce eggs, and genetically determined sexes are still differentiated by phenotype, development, and physiology. Whatever the reason for XY and ZW, us XY'ers at the Raptor Resource Project wish all birds the best of luck this spring!

A list of resources that helped me learn about this topic:
Did you know?