ZW sex-determination system

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ZW sex determination in birds (as exemplified with chickens) Avian sex determination.jpg
ZW sex determination in birds (as exemplified with chickens)

The ZW sex-determination system is a chromosomal system that determines the sex of offspring in birds, some fish and crustaceans such as the giant river prawn, some insects (including butterflies and moths), the schistosome family of flatworms, and some reptiles, e.g. majority of snakes, lacertid lizards and monitors, including Komodo dragons. It is also present in some plants, where it has probably evolved independently on several occasions. [1] The letters Z and W are used to distinguish this system from the XY sex-determination system. In the ZW system, females have a pair of dissimilar ZW chromosomes, and males have two similar ZZ chromosomes.

Contents

In contrast to the XY sex-determination system and the X0 sex-determination system, where the sperm determines the sex, in the ZW system, the ovum determines the sex of the offspring. Males are the homogametic sex (ZZ), while females are the heterogametic sex (ZW). The Z chromosome is larger and has more genes, similarly to the X chromosome in the XY system.

Significance of the ZW and XY systems

No genes are shared between the avian ZW and mammalian XY chromosomes, [2] and, from a comparison between chicken and human, the Z chromosome appears similar to the autosomal chromosome 9 in humans. It has been proposed that the ZW and XY sex determination systems do not share an origin but that the sex chromosomes are derived from autosomal chromosomes of the common ancestor. These autosomes are thought to have evolved sex-determining loci that eventually developed into the respective sex chromosomes once the recombination between the chromosomes (X and Y or Z and W) was suppressed. [3]

The platypus, a monotreme mammal, has a system of five pairs of XY chromosomes. They form a multiple chain due to homologous regions in male meiosis and finally segregates into XXXXX-sperm and YYYYY-sperm. The bird Z-like pair shows up on opposite ends of the chain. Areas homologous to the bird Z chromosome are scattered throughout X3 and X5. [4] : fig. 5  Although the sex-determination system is not necessarily linked to that of birds and definitely not to that of therian mammals, the similarity at least allowed for the conclusion that mammals evolved sex chromosomes twice. [5] The previous report that platypus has X chromosomes similar to that of therian mammals is now considered a mistake. [6]

Bird and snake ZW are unrelated, having evolved from different autosomes. [7] However, the bird-like chromosomes of platypus may indicate that ancestors of snakes had a bird-like ZW system. [6]

Across species

In birds

While there has not been extensive research on other organisms with the ZW sex-determination system, in 2007, researchers announced that chickens' and zebra finches' sex chromosomes do not exhibit any type of chromosome-wide dosage compensation, and instead seem to dosage compensate on a gene-by-gene basis. [8] [9] Specific locations on the chicken Z chromosome, such as the MHM region, are thought to exhibit regional dosage compensation, though researchers have argued that this region does not actually constitute local dosage compensation. [10] [11] Further research expanded the list of birds that do not exhibit any type of chromosome-wide dosage compensation to crows and ratites, thus implying that all avian chromosomes lack chromosome-wide dosage compensation. [12] [13] Both transcriptional and translational gene-specific dosage compensation have been observed in avian sex chromosomes. [14] In addition, the involvement of sex-biased miRNAs was proposed to compensate for the presence of two Z-chromosomes in male birds. [15]

It is unknown whether it might be that the presence of the W chromosome induces female features, or whether instead it is the duplication of the Z chromosome that induces male ones; unlike mammals, no birds with a double W chromosome (ZWW) or a single Z (Z0) have been satisfactorily documented. However, it is known that the removal or damage to the ovaries of female birds can lead to the development of male plumage, suggesting that female hormones repress the expression of male characteristics in birds. [16] It appears possible that either condition could cause embryonic death, or that both chromosomes could be responsible for sex selection. [17] [ better source needed ] One possible gene that could determine sex in birds is the DMRT1 gene. Studies have shown that two copies of the gene are necessary for male sex determination. [14] [18]

The ZW sex-determination system makes it possible to create sex-link chickens in which color at hatching is differentiated by sex, thus making chick-sexing an easier process.

In snakes

Snakes' W chromosomes show different levels of decay compared to their Z chromosomes. This allows for tracking the shrinking of W chromosomes (analogous to the shrinking of Y chromosomes) by comparing across species. Mapping of specific genes reveals that the snake system is different from the bird system. It is not yet known which gene is the sex-determining one in snakes. One thing that stood out was that pythons show little signs of "W-shrinking". [7]

Boa and Python families are now known to probably have an XY sex-determination system. [19] Interest in looking into this came from female family members capable of parthenogenesis, or producing offspring without mating. In 2010 a female Boa constrictor that produced 22 female offspring in this manner was found in the wild. By then it was presumed that such a pattern was produced by WW chromosomes. [20] Python bivittatus and Boa imperator, similarly only produce female offspring; their genomes share male-specific single nucleotide polymorphisms identifiable by restriction enzyme digestion. Their chromosomal origins, however, differ: Python's XY are similar to other snakes' ZW, while Boa XY maps to microchromosomes in other snakes. [21] The female-only pattern is in contrast to the ZW Colubroidean parthenogens, which always produce male (ZZ) offspring. [22]

In moths and butterflies

In Lepidoptera (moths and butterflies), females can have Z, ZZW, or ZZWW. [23]

In schistosomes

The family Schistosomatidae, commonly called blood flukes, are small parasitic flatworms dwelling in the blood vessels of the bladder, liver, intestines and other organs of birds and mammals. They are the only sexually heteromorphic family among the trematode class, and depend on remaining biochemically paired in copula to complete their life cycle. [24] The heterogametic sex chromosomes in females of nine species of schistosomes were first described by geneticist Margaret Menzel and parasitologist Robert B. Short of Florida State University in 1960. [25] [26] The difference in the sex chromosomes was noted during the pachytene stage of meiotic prophase, when the chromosomes thicken and align with their homologous partner.

In turtles

Trionychidae turtles possess a ZZ-ZW sex determinate system, which originated sometime between the beginning of the Jurassic and the Early Cretaceous. [27]

In plants

Among the approximately 5% of plant species that have separate male and female individuals (dioecious), several are known to have a ZW system of sex determination. These include pistachio, several species of strawberry such as Fragaria virginiana and Fragaria chiloensis, and several species of willow including Salix viminalis and Salix purpurea . [1] [28]

See also

Related Research Articles

<span class="mw-page-title-main">Sex</span> Trait that determines an organisms sexually reproductive function

Sex is the trait that determines whether a sexually reproducing organism produces male or female gametes. A male organism produces small mobile gametes, while a female organism produces larger, non-mobile gametes. An organism that produces both types of gamete is called a hermaphrodite. During sexual reproduction, a male and a female gamete fuse to form a zygote, which develops into an offspring that inherits traits from each parent.

<span class="mw-page-title-main">XY sex-determination system</span> Method of determining sex

The XY sex-determination system is a sex-determination system used to classify many mammals, including humans, some insects (Drosophila), some snakes, some fish (guppies), and some plants. In this system, the sex of an individual is determined by a pair of sex chromosomes. In most cases, females have two of the same kind of sex chromosome (XX), and are called the homogametic sex. Males have two different kinds of sex chromosomes (XY), and are called the heterogametic sex.

<span class="mw-page-title-main">Sex-determination system</span> Biological system that determines the development of an organisms sex

A sex-determination system is a biological system that determines the development of sexual characteristics in an organism. Most organisms that create their offspring using sexual reproduction have two common sexes and a few less common intersex variations.

<span class="mw-page-title-main">Y chromosome</span> Sex chromosome in the XY sex-determination system

The Y chromosome is one of two sex chromosomes in therian mammals and other organisms. Along with the X chromosome, it is part of the XY sex-determination system, in which the Y is the sex-determining because it is the presence or absence of Y chromosome that determines the male or female sex of offspring produced in sexual reproduction. In mammals, the Y chromosome contains the SRY gene, which triggers development of male gonads. The Y chromosome is passed only from male parents to male offspring.

<span class="mw-page-title-main">Haldane's rule</span> Observation in evolutionary biology

Haldane's rule is an observation about the early stage of speciation, formulated in 1922 by the British evolutionary biologist J. B. S. Haldane, that states that if — in a species hybrid — only one sex is inviable or sterile, that sex is more likely to be the heterogametic sex. The heterogametic sex is the one with two different sex chromosomes; in therian mammals, for example, this is the male.

<span class="mw-page-title-main">Sexual differentiation</span> Embryonic development of sex differences

Sexual differentiation is the process of development of the sex differences between males and females from an undifferentiated zygote. Sex determination is often distinct from sex differentiation; sex determination is the designation for the development stage towards either male or female, while sex differentiation is the pathway towards the development of the phenotype.

<span class="mw-page-title-main">Sex-determining region Y protein</span> Protein that initiates male sex determination in therian mammals

Sex-determining region Y protein (SRY), or testis-determining factor (TDF), is a DNA-binding protein encoded by the SRY gene that is responsible for the initiation of male sex determination in therian mammals. SRY is an intronless sex-determining gene on the Y chromosome. Mutations in this gene lead to a range of disorders of sex development with varying effects on an individual's phenotype and genotype.

<span class="mw-page-title-main">Sex-chromosome dosage compensation</span>

Dosage compensation is the process by which organisms equalize the expression of genes between members of different biological sexes. Across species, different sexes are often characterized by different types and numbers of sex chromosomes. In order to neutralize the large difference in gene dosage produced by differing numbers of sex chromosomes among the sexes, various evolutionary branches have acquired various methods to equalize gene expression among the sexes. Because sex chromosomes contain different numbers of genes, different species of organisms have developed different mechanisms to cope with this inequality. Replicating the actual gene is impossible; thus organisms instead equalize the expression from each gene. For example, in humans, female (XX) cells randomly silence the transcription of one X chromosome, and transcribe all information from the other, expressed X chromosome. Thus, human females have the same number of expressed X-linked genes per cell as do human males (XY), both sexes having essentially one X chromosome per cell, from which to transcribe and express genes.

<span class="mw-page-title-main">Male</span> Sex of an organism which produces sperm

Male is the sex of an organism that produces the gamete known as sperm, which fuses with the larger female gamete, or ovum, in the process of fertilisation. A male organism cannot reproduce sexually without access to at least one ovum from a female, but some organisms can reproduce both sexually and asexually. Most male mammals, including male humans, have a Y chromosome, which codes for the production of larger amounts of testosterone to develop male reproductive organs.

<span class="mw-page-title-main">Genital ridge</span>

The genital ridge is the precursor to the gonads. The genital ridge initially consists mainly of mesenchyme and cells of underlying mesonephric origin. Once oogonia enter this area they attempt to associate with these somatic cells. Development proceeds and the oogonia become fully surrounded by a layer of cells.

<span class="mw-page-title-main">Heterogametic sex</span> Sex of a species in which the sex chromosomes are not the same

The heterogametic sex is the sex of a species where an individual's gametes have non-matching sex chromosomes. In humans, the heterogametic sex is the male sex, where each gamete's sex chromosomes are X and Y. This is in contrast to the female sex, where each gamete's sex chromosomes are X and X. This arrangement is understood within the XY sex-determination system.

<span class="mw-page-title-main">Sex chromosome</span> Chromosome that differs from an ordinary autosome in form, size, and behavior

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<span class="mw-page-title-main">Parthenogenesis</span> Asexual reproduction without fertilization

Parthenogenesis is a natural form of asexual reproduction in which growth and development of embryos occur in a gamete without combining with another gamete. In animals, parthenogenesis means development of an embryo from an unfertilized egg cell. In plants, parthenogenesis is a component process of apomixis. In algae, parthenogenesis can mean the development of an embryo from either an individual sperm or an individual egg.

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<span class="mw-page-title-main">Female</span> Sex of an organism that produces ova

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<span class="mw-page-title-main">Microchromosome</span> Type of chromosome

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Jennifer Ann Marshall Graves is an Australian geneticist. She is Distinguished Professor within the La Trobe Institute for Molecular Science, La Trobe University, Australia and Professor Emeritus of the Australian National University.

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<span class="mw-page-title-main">Evolution of sex-determining mechanisms</span>

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References

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