Sex determination in Silene

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Silene dioica, female flower Pink flower in Jyvaskyla.jpg
Silene dioica , female flower
Silene dioica male flowers (etamins) with anthers 20160831Silene dioica2.jpg
Silene dioica male flowers (etamins) with anthers
Silene noctiflora, bisexual flower Silene noctiflora sl17.jpg
Silene noctiflora , bisexual flower

Silene is a flowering plant genus that has evolved a dioecious reproductive system. This is made possible through heteromorphic sex chromosomes expressed as XY. [1] [2] Silene recently evolved sex chromosomes 5-10 million years ago and are widely used by geneticists and biologists to study the mechanisms of sex determination since they are one of only 39 species across 14 families of angiosperm that possess sex-determining genes. [3] Silene are studied because of their ability to produce offspring with a plethora of reproductive systems. The common inference drawn from such studies is that the sex of the offspring is determined by the Y chromosome.

Contents

Evolution of sex chromosomes

Biologists have found that sex chromosomes in plants originated from pairs of autosomes. As these chromosomes diverge from their autosomal ancestor and from each other as a homologous pair, they have the potential to increase or decrease in size due to mutation and recombination. In the case of Silene, the pair of automsomal chromosomes are transformed into heteromorphic sex-determining chromosomes expressed as XY. It is important to recognize that not all species of Silene have this sex determination system. A few, such as S. colpophylla, possess homomorphic sex chromosomes. [4]

Plants with sex-determining chromosomes, like Silene, can develop uni-sexual reproductive structures because of the loss and gain of sex-determining genes. Mutations can cause female sterility, male sterility, or adverse combinations of genes that can lead to monoecy, gynodioecy, and dioecy. [4]

Species of Silene with different reproductive systems

The mechanisms involved in the sex determination of Silene are complex and can lead to various reproductive systems among the offspring. The table below provides only a few examples of these possible systems. Those which are most commonly found within this genus are hermaphroditism (monoecious plant with both staminate and pistillate), dioecy (male and female reproductive systems found in separate morphs), and gynodioecy (existence of female and hermaphroditic reproductive systems among the individuals of the population). [ citation needed ]

Sexual systems vary across species most silene species are hermaphroditic representing 58.2% of silene species, 14.3% are dioecious, 13.3%  gynodioecious, and 12.2% being both gynodioecious and gynomonoecious. Trioecy, andromonoecy, and gynomonoecy have also been reported but are extremely rare. [5]

Species Name [6] Reproductive System(s) [6]
S. latifoliadioecy
S. dioicadioecy
S. diclinisdioecy
S. pendulagynodioecy
S. noctifloragynodioecy, hermaphrodite
S. acaulis trioecy
S. saxifragagynodioecy, andromonoecy
S. vulgarisgynodioecy, trioecy
S. gallicahermaphrodite
S. inapertahermaphrodite

Genetics

Out of 300,000 species of angiosperm, Silene are among the 5 to 10 percent whose individual offspring can be of different sexes. [7] [8] Hetermorphic sex-determining chromosomes are very infrequent in plant genera; some notable examples that possess them, other than Silene, are Rumex, Humulus, and Cannabis . [9]

All species of Silene that are diploid possess the same number of chromosomes (n=12); males possess sex-determining Y chromosomes that are much larger than the X chromosomes. [4] [10] Recombination generally occurs only when two X chromosomes (XX) pair during female meiosis. Contrastingly, recombination is suppressed across most of the Y chromosomes during pairing in male meiosis (XY). [11] When recombination does occur in XY chromosomes, it is confined to the tips of the chromosome, leaving most of the genetic material in the Y chromosome intact. [ citation needed ]

Importance of Y-chromosome

Several studies concentrated on Silene latifolia have shown a correlation between loci on the Y chromosome and the sex that is expressed phenotypically in the flower. Two of these sex-linked genes "promote maleness" (male fertility and male promotion) and one of them codes for female suppression. [4] [7] Therefore, the Y chromosome carries three dominant genes that determine the sexual expression in a Silene offspring. If there are deletions and mutations in the Y chromosome during reproduction, then different sex-linked genes are lost. The different combinations of possible sex-related genes that an individual Silene can have in one Y chromosome are what creates the rich variation of sexual phenotypes throughout this genus. [ citation needed ]

This diagram illustrates the different gene combinations that can appear on the Y chromosome of Silene latifolia. Each combination of genes leads to a distinct sexual expression. Adapted from "Silene latifolia : The Classical Model to Study Heteromorphic Sex Chromosomes" by E. Kejnovsky and B. Vyskot. Silene latifolia Y Chromosome.jpg
This diagram illustrates the different gene combinations that can appear on the Y chromosome of Silene latifolia. Each combination of genes leads to a distinct sexual expression. Adapted from "Silene latifolia : The Classical Model to Study Heteromorphic Sex Chromosomes" by E. Kejnovsky and B. Vyskot.

Sexual expression

The system for determining sex in Silene latifolia is close to that found in humans because in both cases the Y chromosome determines what reproductive mechanisms will be expressed in the offspring. They differ because there are multiple sex determining genes on the Y chromosome of S. latifolia, while in humans the presence or absence of the Y chromosome strictly determines whether the offspring is male or female.

Different combinations of the genes present in a Silene Y chromosome affect the sexual expression in the organism. For example, there are two genetic variations that can lead to male sterility in S. latifolia: if it possesses two Y chromosomes (YY) or if the Y chromosome possesses the female suppression and male promoting genes. Silene are also susceptible to a type of sexually transmitted infection that causes sterility. Another possible combination includes the presence of all three sex-linked genes (male suppressing, female suppressing, and male promoting) which produce virile male offspring. The flower is hermaphrodite when both male promoting and male fertility genes are present. Lastly, Y chromosomes carrying both female suppression and male fertility genes creates an asexual organism. [7]

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. 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. By convention, organisms that produce smaller, more mobile gametes are called male, while organisms that produce larger, non-mobile gametes are called female. An organism that produces both types of gamete is hermaphrodite.

<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. 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.

<i>Silene</i> Genus of flowering plants

Silene is a genus of flowering plants in the family Caryophyllaceae. Containing nearly 900 species, it is the largest genus in the family. Common names include campion and catchfly. Many Silene species are widely distributed, particularly in the northern hemisphere.

<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">Sex-chromosome dosage compensation</span> Biological process

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.

Dioecy is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly or indirectly. Dioecious reproduction is biparental reproduction. Dioecy has costs, since only the female part of the population directly produces offspring. It is one method for excluding self-fertilization and promoting allogamy (outcrossing), and thus tends to reduce the expression of recessive deleterious mutations present in a population. Plants have several other methods of preventing self-fertilization including, for example, dichogamy, herkogamy, and self-incompatibility.

<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.

<i>Silene latifolia</i> Species of flowering plant

Silene latifolia, commonly known as white campion, is a dioecious flowering plant in the family Caryophyllaceae, native to most of Europe, Western Asia and northern Africa. It is a herbaceous annual, occasionally biennial or a short-lived perennial plant, growing to between 40–80 centimetres tall. It is also known in the US as bladder campion but should not be confused with Silene vulgaris, which is more generally called bladder campion.

<span class="mw-page-title-main">ZW sex-determination system</span> Chromosomal system

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, 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. 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.

The mechanisms of reproductive isolation are a collection of evolutionary mechanisms, behaviors and physiological processes critical for speciation. They prevent members of different species from producing offspring, or ensure that any offspring are sterile. These barriers maintain the integrity of a species by reducing gene flow between related species.

Cytoplasmic male sterility is total or partial male sterility in hermaphrodite organisms, as the result of specific nuclear and mitochondrial interactions. Male sterility is the failure to produce functional anthers, pollen, or male gametes. Such male sterility in hermaphrodite populations leads to gynodioecious populations.

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

Sex chromosomes are chromosomes that carry the genes that determine the sex of an individual. The human sex chromosomes are a typical pair of mammal allosomes. They differ from autosomes in form, size, and behavior. Whereas autosomes occur in homologous pairs whose members have the same form in a diploid cell, members of an allosome pair may differ from one another.

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

Sexual differentiation in humans is the process of development of sex differences in humans. It is defined as the development of phenotypic structures consequent to the action of hormones produced following gonadal determination. Sexual differentiation includes development of different genitalia and the internal genital tracts and body hair plays a role in sex identification.

Androdioecy is a reproductive system characterized by the coexistence of males and hermaphrodites. Androdioecy is rare in comparison with the other major reproductive systems: dioecy, gynodioecy and hermaphroditism. In animals, androdioecy has been considered a stepping stone in the transition from dioecy to hermaphroditism, and vice versa.

<span class="mw-page-title-main">Hermaphrodite</span> Sexually reproducing organism that produces both male and female gametes

A hermaphrodite is a sexually reproducing organism that produces both male and female gametes. Animal species in which individuals are either male or female are gonochoric, which is the opposite of hermaphroditic.

<span class="mw-page-title-main">Gynodioecy</span> Coexistence of female and hermaphrodite within a population

Gynodioecy is a rare breeding system that is found in certain flowering plant species in which female and hermaphroditic plants coexist within a population. Gynodioecy is the evolutionary intermediate between hermaphroditism and dioecy.

<span class="mw-page-title-main">ZO sex-determination system</span> Biological system in certain moths

The ZO sex-determination system is a system that determines the sex of offspring in several moths. In those species, there is one sex chromosome, Z. Males have two Z chromosomes, whereas females have one Z. Males are ZZ, while females are ZO.

<span class="mw-page-title-main">Sexual system</span> Distribution of male and female functions across a species.

A sexual system is a distribution of male and female function across organisms in a species. The terms reproductive system and mating system have also been used as synonyms.

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