In genetics, transgressive segregation is the formation of extreme phenotypes, or transgressive phenotypes, observed in segregated hybrid populations compared to phenotypes observed in the parental lines. [1] The appearance of these transgressive (extreme) phenotypes can be either positive or negative in terms of fitness. If both parents' favorable alleles come together, it will result in a hybrid having a higher fitness than the two parents. The hybrid species will show more genetic variation and variation in gene expression than their parents. As a result, the hybrid species will have some traits that are transgressive (extreme) in nature. Transgressive segregation can allow a hybrid species to populate different environments/niches in which the parent species do not reside, or compete in the existing environment with the parental species.
Genetic
There are many causes for transgressive segregation in hybrids. One cause can be due to recombination of additive alleles. Recombination results in new pairs of alleles at two or more loci. These different pairs of alleles can give rise to new phenotypes if gene expression has been changed at these loci. Another cause can be elevated mutation rate. When mutation rates are high, it is more probable that a mutation will occur and cause an extreme phenotypic change. Reduced developmental stability is another cause for transgressive segregation. Developmental stability refers to the capability of a genotype to go through a constant development of a phenotype in a certain environmental setting. If there is a disturbance due to genetic or environmental factors, the genotype will be more sensitive to phenotypic changes. Another cause arises from the interaction between two alleles of two different genes, also known as the epistatic effect. Epistasis is the event when one allele at a locus prevents an allele at another locus to express its product as if it is masking its effect. Therefore, epistasis can be related to gene over dominance caused by heterozygosity at specific loci.[2] What this means is that the heterozygote (hybrid) when compared to the homozygote (parent) is better adapted and therefore shows more transgressive, extreme phenotypes. All of these causes lead to the appearance of these extreme phenotypes and creates a hybrid species that will deviate away from the parent species niche and eventually create an individual "hybrid" species.
Environmental
Other than the genetic factors solely causing transgressive segregation, environmental factors can cause genetic factors to take place. Environmental factors that cause transgressive segregation can be influenced by human activity and climate change. Both human activity and climate change have the capability to force species of a specific genome to interact with other species with different genomes.
For example, if a bridge is built that connects two isolated areas to one another, a gene flow door would open. This open door will increase the interactions between different species with different genomes can create hybrid species that can potentially show transgressive phenotypes. Human activity can open the gene flow door by pursuing harmful actions such as cutting down forests and pollution. Climate change on the other hand can open the gene flow door by breaking climate and environmental barriers that were present before. This convergence between species can give rise to a hybrid species that will have more phenotypic variation when compared to the parent species. This increase in phenotypic variation has the potential for transgressive segregation to occur. [2]
In Kenya, there is a fungus called septoria tritici blotch (STB) that diminishes yield in wheat crop. The parent species of wheat had little resistance toward STB, but the hybrid species due to transgressive segregation showed a higher resistance toward STB and therefore a higher fitness. You can create a higher resistance to STB by crossing genes together that are efficient. In result, out of 36 crosses there were 31 that showed a higher mean fitness than the control, parent value. These 31 crosses indicate a higher resistance to STB. The crosses used were from other commercial wheat's that were high yielding which is advantageous because there is a lower chance of deleterious (unwanted traits) appearing and therefore an increase in beneficial traits. Transgressive segregation has been found to be useful to create a resistance toward this organism in order to increase the yield of wheat crop. [3]
Rieseberg used sunflowers to show the transgressive segregation of parental traits. Helianthus annuus and Helianthus petiolaris are the two parent groups for the hybrids. Ultimately there were three hybrid sunflower species. When compared to the fitness of the parents, the hybrids showed a higher tolerance in areas which the parent species would not be able to survive i.e. salt marsh, sand dunes, and deserts. Transgressive segregation allowed these hybrids to survive in areas that the parent would not be able to. Therefore, the hybrids were populated in areas where the parent species were not. This is due to hybrid species showing more gene expression (phenotypes) than their parents and also having some genes that are transgressive (extreme) in nature. [4]
There are many ways to test if transgressive segregation occurred within a population. One common way to test for transgressive segregation is to use a Dunnett's test. This test looks at whether the hybrid species' performance was different from the control group by looking whether or not the mean of the control group (parent species) differs significantly from mean of the other groups. If there is a difference, that is an indication of transgressive segregation. [5] Another commonly used test is the use of quantitative trait loci (QTL) to assess transgressive segregation. Alleles with QTL that were opposed (either by overdomiance or underdominance) of the parental parent QTL indicate that transgressive segregation occurred. Alleles with QTL that was the same as the predicted parent QTL showed that there was no transgressive segregation. [6]
Transgressive segregation creates an opportunity for new hybrid species to arise that are more fit than their ancestors. As seen with the STB in Kenya and Rieseberg's sunflowers, transgressive segregation can be used to create a species that is more adaptable and resistant in areas where there is environmental stress. Transgressive segregation can be seen as genetic engineering in the way that the goal for each of these events is to create an organism that is more fit than the last.
Mendelian inheritance is a type of biological inheritance that follows the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, and popularized by William Bateson. These principles were initially controversial. When Mendel's theories were integrated with the Boveri–Sutton chromosome theory of inheritance by Thomas Hunt Morgan in 1915, they became the core of classical genetics. Ronald Fisher combined these ideas with the theory of natural selection in his 1930 book The Genetical Theory of Natural Selection, putting evolution onto a mathematical footing and forming the basis for population genetics within the modern evolutionary synthesis.
In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and the second recessive. This state of having two different variants of the same gene on each chromosome is originally caused by a mutation in one of the genes, either new or inherited. The terms autosomal dominant or autosomal recessive are used to describe gene variants on non-sex chromosomes (autosomes) and their associated traits, while those on sex chromosomes (allosomes) are termed X-linked dominant, X-linked recessive or Y-linked; these have an inheritance and presentation pattern that depends on the sex of both the parent and the child. Since there is only one copy of the Y chromosome, Y-linked traits cannot be dominant nor recessive. Additionally, there are other forms of dominance such as incomplete dominance, in which a gene variant has a partial effect compared to when it is present on both chromosomes, and co-dominance, in which different variants on each chromosome both show their associated traits.
Genetic linkage is the tendency of DNA sequences that are close together on a chromosome to be inherited together during the meiosis phase of sexual reproduction. Two genetic markers that are physically near to each other are unlikely to be separated onto different chromatids during chromosomal crossover, and are therefore said to be more linked than markers that are far apart. In other words, the nearer two genes are on a chromosome, the lower the chance of recombination between them, and the more likely they are to be inherited together. Markers on different chromosomes are perfectly unlinked.
In population genetics, directional selection, is a mode of negative natural selection in which an extreme phenotype is favored over other phenotypes, causing the allele frequency to shift over time in the direction of that phenotype. Under directional selection, the advantageous allele increases as a consequence of differences in survival and reproduction among different phenotypes. The increases are independent of the dominance of the allele, and even if the allele is recessive, it will eventually become fixed.
A quantitative trait locus (QTL) is a locus that correlates with variation of a quantitative trait in the phenotype of a population of organisms. QTLs are mapped by identifying which molecular markers correlate with an observed trait. This is often an early step in identifying and sequencing the actual genes that cause the trait variation.
Genetic architecture is the underlying genetic basis of a phenotypic trait and its variational properties. Phenotypic variation for quantitative traits is, at the most basic level, the result of the segregation of alleles at quantitative trait loci (QTL). Environmental factors and other external influences can also play a role in phenotypic variation. Genetic architecture is a broad term that can be described for any given individual based on information regarding gene and allele number, the distribution of allelic and mutational effects, and patterns of pleiotropy, dominance, and epistasis.
A polygene is a member of a group of non-epistatic genes that interact additively to influence a phenotypic trait, thus contributing to multiple-gene inheritance, a type of non-Mendelian inheritance, as opposed to single-gene inheritance, which is the core notion of Mendelian inheritance. The term "monozygous" is usually used to refer to a hypothetical gene as it is often difficult to characterise the effect of an individual gene from the effects of other genes and the environment on a particular phenotype. Advances in statistical methodology and high throughput sequencing are, however, allowing researchers to locate candidate genes for the trait. In the case that such a gene is identified, it is referred to as a quantitative trait locus (QTL). These genes are generally pleiotropic as well. The genes that contribute to type 2 diabetes are thought to be mostly polygenes. In July 2016, scientists reported identifying a set of 355 genes from the last universal common ancestor (LUCA) of all organisms living on Earth.
Inbreeding depression is the reduced biological fitness in a given population as a result of inbreeding. Population biological fitness refers to an organism's ability to survive and perpetuate its genetic material. Inbreeding depression is often the result of a population bottleneck. In general, the higher the genetic variation or gene pool within a breeding population, the less likely it is to suffer from inbreeding depression.
Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Introgression is a long-term process, even when artificial; it may take many hybrid generations before significant backcrossing occurs. This process is distinct from most forms of gene flow in that it occurs between two populations of different species, rather than two populations of the same species.
In biology, a cline is a measurable gradient in a single character of a species across its geographical range. First coined by Julian Huxley in 1938, the “character” of the cline referred to is usually genetic, or phenotypic. Clines can show smooth, continuous gradation in a character, or they may show more abrupt changes in the trait from one geographic region to the next.
Marker assisted selection or marker aided selection (MAS) is an indirect selection process where a trait of interest is selected based on a marker linked to a trait of interest, rather than on the trait itself. This process has been extensively researched and proposed for plant and animal breeding.
A doubled haploid (DH) is a genotype formed when haploid cells undergo chromosome doubling. Artificial production of doubled haploids is important in plant breeding.
In genetics, association mapping, also known as "linkage disequilibrium mapping", is a method of mapping quantitative trait loci (QTLs) that takes advantage of historic linkage disequilibrium to link phenotypes to genotypes, uncovering genetic associations.
In statistical genetics, inclusive composite interval mapping (ICIM) has been proposed as an approach to QTL mapping for populations derived from bi-parental crosses. QTL mapping is based on genetic linkage map and phenotypic data and attempts to locate individual genetic factors on chromosomes and to estimate their genetic effects.
Quantitative trait loci mapping or QTL mapping is the process of identifying genomic regions that potentially contain genes responsible for important economic, health or environmental characters. Mapping QTLs is an important activity that plant breeders and geneticists routinely use to associate potential causal genes with phenotypes of interest. Family-based QTL mapping is a variant of QTL mapping where multiple-families are used.
Linkage based QTL mapping is a variant of QTL mapping.
Phenotypic Integration is a metric for measuring the correlation of multiple functionally-related traits to each other. Complex phenotypes often require multiple traits working together in order to function properly. Phenotypic integration is significant because it provides an explanation as to how phenotypes are sustained by relationships between traits. Every organism's phenotype is integrated, organized, and a functional whole. Integration is also associated with functional modules. Modules are complex character units that are tightly associated, such as a flower. It is hypothesized that organisms with high correlations between traits in a module have the most efficient functions. The fitness of a particular value for one phenotypic trait frequently depends on the value of the other phenotypic traits, making it important for those traits evolve together. One trait can have a direct effect on fitness, and it has been shown that the correlations among traits can also change fitness, causing these correlations to be adaptive, rather than solely genetic. Integration can be involved in multiple aspects of life, not just at the genetic level, but during development, or simply at a functional level. Integration can be caused by genetic, developmental, environmental, or physiological relationships among characters. Environmental conditions can alter or cause integration, i.e. they may be plastic. Correlational selection, a form of natural selection can also produce integration. At the genetic level, integration can be caused by pleiotropy, close linkage, or linkage disequilibrium among unlinked genes. At the developmental level it can be due to cell-cell signaling such as in the development of the ectopic eyes in Drosophila. It is believed that the patterns of genetic covariance helped distinguish certain species. It can create variation among certain phenotypes, and can facilitate efficiency. This is significant because integration may play a huge role in phenotypic evolution. Phenotypic integration and its evolution can not only create large amounts of variety among phenotypes which can cause variation among species. For example, the color patterns on Garter snakes range widely and are caused by the covariance among multiple phenotypes.
Genetic variance is a concept outlined by the English biologist and statistician Ronald Fisher in his fundamental theorem of natural selection. In his 1930 book The Genetical Theory of Natural Selection, Fisher postulates that the rate of change of biological fitness can be calculated by the genetic variance of the fitness itself. Fisher tried to give a statistical formula about how the change of fitness in a population can be attributed to changes in the allele frequency. Fisher made no restrictive assumptions in his formula concerning fitness parameters, mate choices or the number of alleles and loci involved.
This glossary of evolutionary biology is a list of definitions of terms and concepts used in the study of evolutionary biology, population biology, speciation, and phylogenetics, as well as sub-disciplines and related fields. For additional terms from related glossaries, see Glossary of genetics, Glossary of ecology, and Glossary of biology.
Eukaryote hybrid genomes result from interspecific hybridization, where closely related species mate and produce offspring with admixed genomes. The advent of large-scale genomic sequencing has shown that hybridization is common, and that it may represent an important source of novel variation. Although most interspecific hybrids are sterile or less fit than their parents, some may survive and reproduce, enabling the transfer of adaptive variants across the species boundary, and even result in the formation of novel evolutionary lineages. There are two main variants of hybrid species genomes: allopolyploid, which have one full chromosome set from each parent species, and homoploid, which are a mosaic of the parent species genomes with no increase in chromosome number.