In addition to the normal karyotype, wild populations of many animal, plant, and fungi species contain B chromosomes (also known as supernumerary,accessory, (conditionally-)dispensable, or lineage-specific chromosomes). [1] By definition, these chromosomes are not essential for the life of a species, and are lacking in some (usually most) of the individuals. Thus a population would consist of individuals with 0, 1, 2, 3 (etc.) B chromosomes. [1] B chromosomes are distinct from marker chromosomes or additional copies of normal chromosomes as they occur in trisomies.
The evolutionary origin of supernumerary chromosomes is obscure, but presumably, they must have been derived from heterochromatic segments of normal chromosomes in the remote past. In general "we may regard supernumeraries as a very special category of genetic polymorphism which, because of manifold types of accumulation mechanisms, does not obey the ordinary Mendelian laws of inheritance." [1]
Next generation sequencing has shown that the B chromosomes from rye are amalgamations of the rye A chromosomes. [2] Similarly, B chromosomes of the cichlid fish Haplochromis latifasciatus also have been shown to arise from rearrangements of normal A chromosomes. [3]
Most B chromosomes are mainly or entirely heterochromatic (i.e. largely non-coding), but some contain sizeable euchromatic segments [4] (e.g. such as the B chromosomes of maize). In some cases, B chromosomes act as selfish genetic elements. In other cases, B chromosomes provide some positive adaptive advantage. For instance, the British grasshopper Myrmeleotettix maculatus has two structural types of B chromosomes: metacentrics and submetacentric. The supernumeraries, which have a satellite DNA, occur in warm, dry environments, and are scarce or absent in humid, cooler localities.
There is evidence of deleterious effects of supernumeraries on pollen fertility, and favourable effects or associations with particular habitats are also known in a number of species.[ citation needed ]
B chromosomes have a tendency to accumulate in meiotic cell products resulting in an increase of B number over generations, thereby acting as selfish genetic elements. However, this effect is counterbalanced for selection against infertility.
Chromosome polymorphisms are very common among fungi. Different isolates of the same species often have a different chromosome number, with some of these additional chromosomes being unnecessary for normal growth in culture. The extra chromosomes are known as conditionally dispensable, or supernumerary, because they are dispensable for certain situations, but may confer a selective advantage under different environments. [5]
Supernumerary chromosomes do not carry genes that are necessary for basic fungal growth but may have some functional significance. For example, it has been discovered that the supernumerary chromosome of the pea pathogen Haematonectria haematococca carries genes that are important to the disease-causing capacity of the fungus. This supernumerary DNA was found to code for a group of enzymes that metabolize toxins, known as phytoalexins, that are secreted by the plant's immune system. [5] It is possible that these supernumerary elements originated in horizontal gene transfer events because sequence analysis often indicates that they have a different evolutionary history from essential chromosomal DNA. [5]
The wheat-infecting fungal pathogen Zymoseptoria tritici contains 8 dispensable B-chromosomes, which is the largest number of dispensable chromosomes observed in fungi. [6]
B-chromosomes are a significant reflection of genetic diversity between varying plant species. [7] These supernumerary chromosomes are commonly observed in angiosperms, specifically the flowering plants bred through outcrossing. [7]
The obscure development of B-chromosomes is supported by the irregularity of their appearances in specific species populations. The number of B-chromosomes copied between cells within individual members of a plant population fluctuates. [8] For example, the sister species Aegilops speltoides and Aegilops mutica possess copies of B-chromosomes within their aerial tissues, while their roots exhibit an absence of these supernumerary chromosomes. [8]
The morphological structure and size of B-chromosomes is different from normally-occurring chromosomes in both plants and mammals. [8] Most often in plants, B-chromosomes are notably “non-homologous and smaller than the smallest A-chromosome”. [8]
In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA. The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as regulatory sequences, and often a substantial fraction of junk DNA with no evident function. Almost all eukaryotes have mitochondria and a small mitochondrial genome. Algae and plants also contain chloroplasts with a chloroplast genome.
Ploidy is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively, in each homologous chromosome pair, which chromosomes naturally exist as. Somatic cells, tissues, and individual organisms can be described according to the number of sets of chromosomes present : monoploid, diploid, triploid, tetraploid, pentaploid, hexaploid, heptaploid or septaploid, etc. The generic term polyploid is often used to describe cells with three or more sets of chromosomes.
Selfish genetic elements are genetic segments that can enhance their own transmission at the expense of other genes in the genome, even if this has no positive or a net negative effect on organismal fitness. Genomes have traditionally been viewed as cohesive units, with genes acting together to improve the fitness of the organism. However, when genes have some control over their own transmission, the rules can change, and so just like all social groups, genomes are vulnerable to selfish behaviour by their parts.
Polyploidy is a condition in which the cells of an organism have more than one pair of (homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two complete sets of chromosomes, one from each of two parents; each set contains the same number of chromosomes, and the chromosomes are joined in pairs of homologous chromosomes. However, some organisms are polyploid. Polyploidy is especially common in plants. Most eukaryotes have diploid somatic cells, but produce haploid gametes by meiosis. A monoploid has only one set of chromosomes, and the term is usually only applied to cells or organisms that are normally diploid. Males of bees and other Hymenoptera, for example, are monoploid. Unlike animals, plants and multicellular algae have life cycles with two alternating multicellular generations. The gametophyte generation is haploid, and produces gametes by mitosis; the sporophyte generation is diploid and produces spores by meiosis.
Genetic recombination is the exchange of genetic material between different organisms which leads to production of offspring with combinations of traits that differ from those found in either parent. In eukaryotes, genetic recombination during meiosis can lead to a novel set of genetic information that can be further passed on from parents to offspring. Most recombination occurs naturally and can be classified into two types: (1) interchromosomal recombination, occurring through independent assortment of alleles whose loci are on different but homologous chromosomes ; & (2) intrachromosomal recombination, occurring through crossing over.
In molecular biology and genetics, transformation is the genetic alteration of a cell resulting from the direct uptake and incorporation of exogenous genetic material from its surroundings through the cell membrane(s). For transformation to take place, the recipient bacterium must be in a state of competence, which might occur in nature as a time-limited response to environmental conditions such as starvation and cell density, and may also be induced in a laboratory.
Agrobacterium is a genus of Gram-negative bacteria established by H. J. Conn that uses horizontal gene transfer to cause tumors in plants. Agrobacterium tumefaciens is the most commonly studied species in this genus. Agrobacterium is well known for its ability to transfer DNA between itself and plants, and for this reason it has become an important tool for genetic engineering.
Powdery mildew is a fungal disease that affects a wide range of plants. Powdery mildew diseases are caused by many different species of ascomycete fungi in the order Erysiphales. Powdery mildew is one of the easier plant diseases to identify, as the signs of the causal pathogen are quite distinctive. Infected plants display white powdery spots on the leaves and stems. This mycelial layer may quickly spread to cover all of the leaves. The lower leaves are the most affected, but the mildew can appear on any above-ground part of the plant. As the disease progresses, the spots get larger and denser as large numbers of asexual spores are formed, and the mildew may spread up and down the length of the plant.
In biological taxonomy, race is an informal rank in the taxonomic hierarchy for which various definitions exist. Sometimes it is used to denote a level below that of subspecies, while at other times it is used as a synonym for subspecies. It has been used as a higher rank than strain, with several strains making up one race. Races may be genetically distinct populations of individuals within the same species, or they may be defined in other ways, e.g. geographically, or physiologically. Genetic isolation between races is not complete, but genetic differences may have accumulated that are not (yet) sufficient to separate species.
Intragenomic conflict refers to the evolutionary phenomenon where genes have phenotypic effects that promote their own transmission in detriment of the transmission of other genes that reside in the same genome. The selfish gene theory postulates that natural selection will increase the frequency of those genes whose phenotypic effects cause their transmission to new organisms, and most genes achieve this by cooperating with other genes in the same genome to build an organism capable of reproducing and/or helping kin to reproduce. The assumption of the prevalence of intragenomic cooperation underlies the organism-centered concept of inclusive fitness. However, conflict among genes in the same genome may arise both in events related to reproduction and altruism.
C-value is the amount, in picograms, of DNA contained within a haploid nucleus or one half the amount in a diploid somatic cell of a eukaryotic organism. In some cases, the terms C-value and genome size are used interchangeably; however, in polyploids the C-value may represent two or more genomes contained within the same nucleus. Greilhuber et al. have suggested some new layers of terminology and associated abbreviations to clarify this issue, but these somewhat complex additions are yet to be used by other authors.
Genome size is the total amount of DNA contained within one copy of a single complete genome. It is typically measured in terms of mass in picograms or less frequently in daltons, or as the total number of nucleotide base pairs, usually in megabases. One picogram is equal to 978 megabases. In diploid organisms, genome size is often used interchangeably with the term C-value.
Common wheat, also known as bread wheat, is a cultivated wheat species. About 95% of wheat produced worldwide is common wheat; it is the most widely grown of all crops and the cereal with the highest monetary yield.
Blumeria graminis is a fungus that causes powdery mildew on grasses, including cereals. It is the only species in the genus Blumeria. It has also been called Erysiphe graminis and Oidium monilioides or Oidium tritici.
Zymoseptoria tritici, synonyms Septoria tritici, Mycosphaerella graminicola, is a species of filamentous fungus, an ascomycete in the family Mycosphaerellaceae. It is a wheat plant pathogen causing septoria leaf blotch that is difficult to control due to resistance to multiple fungicides. The pathogen today causes one of the most important diseases of wheat.
In biology, a pathogen, in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a germ.
Plant–fungus horizontal gene transfer is the movement of genetic material between individuals in the plant and fungus kingdoms. Horizontal gene transfer is universal in fungi, viruses, bacteria, and other eukaryotes. Horizontal gene transfer research often focuses on prokaryotes because of the abundant sequence data from diverse lineages, and because it is assumed not to play a significant role in eukaryotes.
Drosophila neotestacea is a member of the testacea species group of Drosophila. Testacea species are specialist fruit flies that breed on the fruiting bodies of mushrooms. These flies will choose to breed on psychoactive mushrooms such as the Fly Agaric Amanita muscaria. Drosophila neotestacea can be found in temperate regions of North America, ranging from the north eastern United States to western Canada.
Fungal genomes are among the smallest genomes of eukaryotes. The sizes of fungal genomes range from less than 10 Mbp to hundreds of Mbp. The average genome size is approximately 37 Mbp in Ascomycota, 47 Mbp in Basidiomycota and 75 Mbp in Oomycota. The sizes and gene numbers of the smallest genomes of free-living fungi such as those of Wallemia ichthyophaga, Wallemia mellicola or Malassezia restricta are comparable to bacterial genomes. The genome of the extensively researched yeast Saccharomyces cerevisiae contains approximately 12 Mbp and was the first completely sequenced eukaryotic genome. Due to their compact size fungal genomes can be sequenced with less resources than most other eukaryotic genomes and are thus important models for research. Some fungi exist as stable haploid, diploid, or polyploid cells, others change ploidy in response to environmental conditions and aneuploidy is also observed in novel environments or during periods of stress.