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Terminologies in Genetics and Plant Breeding Part -10

M. A general term for mutagenic treatment. M0 refers to the parent material likely to bemutagenized. M1, M2, M3,…are symbols used to designate first, second, third, etc.generations after treatment with a mutagenic agent. M2 is often assumed to beanalogous to F2 generation so far as segregation events and relative degree of geneticvariability is concerned. However, M1 cannot be compared to F1; while there is 100%heterozygosity in F1, mutant loci in M1 are random and not so frequent, hence,heterozygosity is variable.

Macro restriction map: Map depicting the order of and distance between sitesat which restriction enzymes cleave chromosomes.

Macro-evolution. Evolution above species level.

Macromolecule. A large molecule with colloidal properties; a long polymer composed ofmonomers. The molecular weight of such a molecule is greater than 1000. Polymerssuch as DNA, RNA, a protein or a polysaccharide are macromolecules.

Macro-mutation. A mutation with the recognizable effect on a single plant.

Macromutation: a mutation that results in a profound change in an organism,such as a change in a regulatory gene that controls the expression of manystructural genes, as opposed to micromutation

Macrophytes. Rooted or large floating plants generally growing in shallow water only(as in a pond).Maisinte. A maize-teosinte hybrid fodder crop. It has been produced by backcrossingmaize-teosinte hybrid with maize. Its main features are quicker growth, earliermaturity, higher fodder yield and high crude protein content than parents. Owing toteosinte genes, it may give good response to stress conditions.

Main effect QTLA QTL that produces direct effect on expression of the concerned trait.

Maintainer: Used for maintaining and multiplication of a cytoplasmic malesterile line; usually genotypes containing the normal cytoplasm and recessive atthe restorer locus

Major Genes. Genes with large, easily recognizable and relatively stable effects. Thereare numerous examples of well defined morphological or physiological characterswhich are governed by single genes, and are little affected by either the genetic orphysical environments, e.g., differences between field and sweet corn, indeterminatevs. determinate growth habit in fenugreek, etc. It is, however, interesting to mentionthat genes are neither major nor minor; nor are they qualitative/quantitative; rather itis their effects which appear as either major or minor. However, the term such asmajor genes is used frequently, and it is accepted usage.

Major QTLA main effect QTL that explains 10 % or more of the phenotypic variance for the concerned trait.

Male gametocide: Any substance that kills the male reproductive cells of aplant (pollen or pollen mother cells), rendering it male-sterile

Male Sterility. Absence or non-functioning of pollens in plants. This provides a barrierto self-fertilization. However, it is advantageous in hybrids seed production since it138makes mechanical emasculation unnecessary. It occurs spontaneously in almost allcrop plants irrespective of their pollination behaviour as a consequence of mutation atany one of the several loci, which condition vital steps in the formation of functionalpollen.

Male sterility: Producing no functional pollen

Malthusian. Relating to Thomas Robert Malthus and his population theory (1798). Thetheory states that population numbers tend to increase at a faster rate (geometricprogression) than means of subsistence (arithmetic progression), leading tocompetition for environmental resources in short supply.

Map distance: The distance between any two markers on a genetic map, basedon the percentage of crossing-over; the minimum distance between linked genesis 1% and maximum 50%

Map Unit. A unit of distance in a linkage map. It is the “distance” between two linkedgene pairs, recombination value for which is one per cent.

Map-based cloning: The isolation of important genes by cloning the genein question on the basis of molecular maps, where the biochemical function isunknown; it involves the identification of a mutant phenotype for the trait ofinterest (obtained by mutagenesis or from natural variation) and genetic finemapping using many progeny plants; this map is then used for chromosomewalking or landing, with the help of large-insert DNA libraries or physical mapsto isolate the gene

Mapping Function. The relationship between distance in a linkage map andrecombination value (between the two linked genes) or frequency. It is expressed by aformula: true map units = -ln [(1-2 R.F.)/2] x 100, whereas, R.F. stands forrecombination frequency.

Mapping functionsFormulas used for converting recombination frequency into genetic distance.

Mapping populationA population that is suitable for linkage mapping of genetic markers, genes, and/or QTLs.

Mapping. The linear arrangement of genes on a chromosome.

Mapping: The process of identifying the location of a gene or DNA segmentalong a chromosome. In genetic mapping, this is done by analysing patterns ofinheritance in segregating populations (measured in recombinational units,commonly centimorgans). Inphysical mapping, this describes the actual location ofa sequence in a particular genomic region (measured in bp)

Marcotting Technique. A technique to induce rooting of nodes by covering themselectively with suitable materials such as mud, soil-coconut fibre mixture or peatmixtures. The rooted nodes are then cut and replanted in the soil. This facilitatesreduction in the height of the prospective parents to be crossed (as in sugar cane).

Marker Assisted Back Crossing (MABC). Introgression of specific trait(s) from adonor parent into the genetic background of a recurrent parent (generally leadingvariety) using molecular markers (Hospital 2005). The product of MABC is aline/cultivar containing only the major gene/QTL from the donor parent, whileretaining the whole genome of the recurrent parent. MABC approach generallyinvolves transfer of a limited number of trait loci including transgenes from onegenetic background (donor genotype) to the other genetic background (elite variety).MABC approach can also be used to generate near-isogenic lines (NILs) orchromosome segment substitution lines (CSSLs) for genomics research, which arepopulations that are often used for genetic analysis of genes/QTLs. Gene pyramidingis an important application of MABC in which a few different genes for the same trait139(e.g., resistance to different races) or for different traits are brought together in onegenetic background using molecular markers.

Marker indexThe product of multiplex ratio and the average PIC score for a marker system in a given population.

Marker-assisted backcrossingA backcross program based on molecular markers.

Marker-assisted introgression: The use of DNA markers to increase the speedand efficiency of introgression of a new gene or genes into a population; themarkers will be closely linked to the genes in question

Marker-assisted plant breedingThe use of molecular marker data for enhancing the effectiveness of various breeding activities, including planning and execution of breeding programs, and improving selection efficiency.

Marker-Assisted Recurrent Selection (MARS). Estimation of marker effects fromgenotyping F2 or F3 population and phenotyping F2 derived F4 or F5 progenies,followed by two or three recombination cycles based on presence of marker alleles forsmall effect QTLs (Eathington et al. 2007). In the first step of MARS, de novo QTLidentification is carried out initially, that is, QTLs are identified in the breedingpopulation itself, generally derived from good × good crosses. Subsequently, the linescarrying superior alleles for maximum QTLs are crossed to pyramid superior allelesin one genetic background. Recombined lines are then subjected to a final phenotypicscreening to select the best lines for multi-location field traits to release them asvarieties. MARS is particularly useful for capturing the several genomic regionsespecially to target more number of minor as well as major QTLs. Therefore, geneticgain achieved by MARS as compared to the MABC programme is higher (Bernardoand Charcosset 2006). The recurrent-selection method is routinely used mainly incross-pollinated crops like maize, and this process can be improved with the help ofmolecular markers; therefore, the process is called marker-assisted recurrent selectionor MARS.

Marker-Assisted Selection (MAS). The selection of target gene(s) aided by tightlylinked markers. The essential requirements for MAS in a breeding programme are: (a)co-segregation or very close linkage (<1cM) of markers with the target gene/ trait, (b)availability of an efficient means of screening large populations for molecularmarkers, and (c) high reproducibility across laboratories of the screening techniques.Besides, markers and screening techniques should be economical and user-friendly.Of course, the population can be screened at any growth stage and in anyenvironment. It is also independent of interaction effects. Markers can also provideadditional information on the breeding value of a genotype, and thus accelerateselection and breeding cycles. It can be practiced especially for traits whosephenotypic selection is difficult. It may be of special value in breeding species thathave large stature or long generation time (arbour crops) where fewer individuals140might save several hectares, and fewer generations may save several decades. MAS isbeing used efficiently: (a) in gene pyramiding, (b) in marker-assisted alien geneintrogression, and (c) for simultaneous identification and pyramiding of QTLs fromprimitive cultivars and alien species.

Marker-assisted selection (MAS): Use of genetic/DNA markers to guide inchoosing specific plants or lines with targeted traits for new variety development.

Marker-assisted selectionSelection for the desirable allele of a gene/quantitative trait locus (QTL) on the basis of molecular marker(s) linked to it in place of the phenotype generated by this allele.

Markers (Morphological). Traits or genes (variant alleles), which are used to label abiological system throughout the course of experimentation. These are usually mutantalleles, which may be either dominant or recessive. These are limited in number,influenced by environments, and developmental stage specific compared to molecularmarkers. Further, they could appear as pleiotropic effect of other major genes.

Masking Action. Gene action such that a gene hides the effect of another non-allelicgene (when both are present). In oats, for instance, Y produces yellow seed coat and Bproduces black seed coat. The gene Y will have no effect in presence of B, since blackcolour masks yellow ones. It gives a ratio of 12 black (B-Y-, B-yy): 3 yellow (bbY-): 1white (bbyy).

Mass Pedigree Method. A system of breeding in which a genetically diverse populationis propagated in mass until conditions favourable for artificial selection occur, afterwhich pedigree selection is practised. The bulking may end as early as F2 generationor may be continued for many generations if the advent of conditions suitable forselection is long-delayed.

Mass pedigree selection: A system of breeding in which a population ispropagated in bulk until conditions favorable for selection occur; usually, aftermass pedigree selection, pedigree selection follows

Mass Selection. A form of selection in which individual plants are selected dependingupon their better phenotypic values (usually without progeny test) and the nextgeneration is propagated from aggregate of their seeds. In autogamous crop species, ithas basically two uses: First, it can effect improvement in land varieties very safelyand rapidly, and the second, it can be used to purify existing varieties. In the crosspollinatedspecies, it is used to improve characters having high heritability. It isvirtually powerless to effect any improvement in a trait like yield (low heritability)obviously for three reasons: (a) selection is exclusively based on phenotype, (b) thereis lack of pollen control on female parent, and (c) there is no progeny test. Therefore,simplicity of mass selection is the weakness in itself. However, some form of massselection is almost always practiced in most breeding methods.

Mass selection: A form of breeding in which individual plants are selectedon their individual advantages and the next generation propagated from theaggregate of their seeds; the easiest method is to select and multiply together thoseindividuals from a mixture of phenotypes, which correspond to the breeding aim(positive mass selection); it is still applied in cross-pollinating of vegetable species,such as carrots, radishes, or beetroots, in order to improve the uniformity; whenall undesired off-types are rouged in grown crop population, and the remainingindividuals are propagated further, the method is termed negative mass selection;negative mass selection is no longer an adequate breeding method for highlyadvanced varieties; it is usually applied in multiplication of established varieties(i.e., for seed production in order to remove diseased plants, casual hybrids, orother defects)

Maternal Effect. The environmental influence of the mother’s tissues on the phenotypeof the offspring. It may be result of nuclear genes as observed for cases like shellcoilingexample in the water snail Limnaea.

Maternal effect: Any nonlasting environmental effect or influence of thematernal genotype or phenotype on the immediate offspring

Maternal Inheritance. A type of uniparental inheritance in which all progeny have thegenotype and phenotype of the parent acting as the female. It was first discovered byCarl Correns (1909) in his studies on four-o’clock plants (Mirabilis jalapa) for leafvariegation (patches of green and white tissues in the leaf).

Maternal inheritance: Phenotypic differences found between individuals ofidentical genotype due to an effect of maternal inheritance

Maternally Expressed Gene. A gene that contributes to the phenotype of an offspring onthe basis of its expression in the mother.

Mating System/ Mating. Any of a number of schemes by which individuals are assortedin pairs leading to sexual reproduction, and thus formation of zygotes.

Mating Types. The analogous terms used in lower organisms for sexes in higherorganisms. Mating types differ only physiologically, and not in the physical form.

Matriclinous Inheritance. A type of inheritance in which all offspring have the nucleusdeterminedphenotype of the mother.

Matriclinous. Having predominantly maternal hereditary traits.

Matrix substanceAn organic molecule that has the same energy absorption spectrum as the selected laser wavelength and does not interact chemically with the target biomolecule.

Maximum Likelihood. A statistical procedure for estimating values of populationparameters from sample data. The method identifies values that have maximumprobability of being the best fitting for any given set of observations.

Mean Deviation. The sum total of average deviations (d) of individual observationsfrom their arithmetic mean without regard to the sign {􀂦d/N}. Though not usedfrequently, it finds its greatest application in calculating combining ability effect(s) ofa given parent of/or a cross in plant breeding experiments.

Mean. Also called arithmetic mean. It is the arithmetic average and is obtained when thenumber of individuals in the data divides the sum of the values of the observations (inthat data). It is usually designated by μ and x in the population and sample,respectively. Unlike median and mode, it is affected by the extreme values. However,it is most commonly used and preferred as it provides more information about thecentral tendency than other measures.

Mechanical Errors. Errors occurring in the execution of an experiment. These can occurduring layout, management and care of the experiment and data keeping andrecording. These are primarily human errors, and thus cannot be effectively controlledby statistical techniques. These can make a substantial difference in the experimentalresults. Their prevention depends largely on the awareness and the skill of theresearcher in relation to the various operations of the experiment.

Median. The middle value of a series of observations arranged in ascending ordescending order of their magnitudes. It divides the series into two equal halves, halfthe number of the observations lying above it and half below. If the number ofobservations in a series is odd, then (N+1)/2 is the middle item of the series (where Nis the total number of observations). However, in case of an even series data, twomiddle items [(N/2) and (N/2+1)] will represent the median.

Medium. Any material on which experimental cultures are grown. Although compositionof a medium varies according to the objectives of a programme, it usually consists ofsugar (as a source of energy), certain inorganic salts, one or a few phytohormones andvitamins, and agar (as a solidifying agent if a solid medium is to be prepared).

Mega Base Pairs. One million nucleotide pairs.

Megabase (Mb): Unit of length for DNA fragments equal to 1 millionnucleotides and roughly equal to 1 cM.

Megaenvironment. Broad (not necessarily continuous often transcontinental) area withsimilar biotic and abiotic stresses, cropping systems and consumer preferences.Analysis of megaenvironment is performed by AMMI model.

Megagametogenesis: The development of the female gametophyte from afunctional megaspore

Megasporangium. A sporangium producing megaspores.

Megaspore Mother Cell. The diploid (2n) cell in the ovary that gives rise to four haploidmegaspores through meiosis.Meiocyte. A cell embarking upon meiosis (or a cell in which meiosis takes place).

Megaspore. One of the four haploid spores originating from the meiotic division of thediploid megaspore mother cell in the ovary, and which gives rise to the megagametophyte.

Megasporogenesis: The development of the megaspore from the archesporial cell

Meiocyte: The sporocyte giving rise to the embryo sac and to pollen grains

Meiosis. A kind of nuclear division occurring in reproductive cells, which results inproduction of gametes (animals) or sexual spores (in plants) with haploid (n)143chromosome number. Thus, it maintains chromosome number of sexuallyreproducing species.

Meiosis: A type of nuclear division that occurs at some stage in the life cycleof sexually reproducing organisms; by a specific mechanism, the number ofchromosomes is halved to prevent doubling in each generation

Meiospores. Products of meiosis in plants.

Meiotic Drive. Any mechanism that affects the genetic composition of a populationthrough unequal contribution of gene(s) [and thus chromosome(s)] to the gametesduring meiosis. This preferential segregation makes a disproportionate contribution tothe gene pool of the next generation. It is an exception to the rule of 1:1 recovery ofsegregating alleles in heterozygous organism. This unequal or preferential segregationhas also been termed segregation distortion.

Melting Pot Technique. A modified polycross method as in sugar cane. It consists ofplacing arrows from a number of prospective parents in small containers (added withpreservative solution) and effecting random intermating by changing the position ofcontainers (along with arrows) in relation to each other every day. Seeds so obtainedinclude selfs and all possible crosses including reciprocals.

Melting temperature (Tm)The temperature at which 50 % of the DNA duplexes would dissociate into separate strands.

Melting. Denaturation of DNA.

Melting: Denaturation of DNA.

Mendel ’s laws of inheritance: The inheritance of chromosomal genes onthe basis of the chromosome theory of heredity; three laws are considered: (1)law of dominance or of uniformity of hybrids, (2) law of segregation, (3) law ofindependent assortment

Mendelian Inheritance. Inheritance of chromosomal genes in contrast to extrachromosomalinheritance, which deals with cytogenes.

Mendelian population: A group of (potentially) interbreeding plants (crossfertilizingcrops), which may occur in a certain area or in an experimental design

Mendelian Ratio. A ratio of progeny phenotypes based on Mendel’s laws of inheritance.

Mendelian ratio: The segregation ratios according to Mendel ’s laws ofinheritance

Mendelian Variation. The diverse types resulting from gene mutations as well asrecombinations following hybridization between types carrying these mutations. Thisis the major kind of variation exploited by plant breeders.

Meristem Tip Culture. The in vitro culturing of plant tissue from the meristem tipregion for the purpose of regenerating pathogen-free plants.

Meristem. An area of rapidly dividing plant cells. It may be a single cell (as in lowerplants) or it may include many cells (as in higher plants).

Mesolithic Age. The cultural period between the Paleolithic and Neolithic Ages;appearance of the bow and cutting tools.Metabolism. The chemical reactions occurring in a living cell.

Messenger DNA (mDNA): A single-stranded DNA that acts as a messenger ofthe protein biosynthesis

Messenger RNA (mRNA): A single-stranded RNA molecule responsible for the transmission to the ribosomes of the genetic information contained in the nuclear DNA; it is synthesized during transcription and its base sequencing exactly matches that of one of the strands of the double-stranded DNA molecule

Meta-analysisCombining the results from many different studies concerning a single research issue to identify common patterns, sources of disagreements, and any other relationships among their findings.

Metabolic QTLsThese QTLs control the rates of various metabolic reactions and metabolite levels.

Metabolite. A product of metabolism.

MetabolomeAll the metabolites, representing the end products of cellular processes, present in a cell, tissue, organ, or organism.

Metabolomics: An extended discipline of biochemistry that involves the analysis (usually high throughput or broad scale) of small-molecule metabolites and polymers, such as starch; it involves descriptions of biological pathways andcurrent metabolomic databases, such as the Kyoto Encyclopedia of Genes andGenomes (KEGG).

MetabolomicsA systematic study of the characteristic small-molecule metabolite profiles generated by the various cellular metabolic processes.

Metacentric Chromosome. A chromosome with centromere placed in the middle so thatits both arms are equal, and take V-shape at anaphase.

Metacentric: Applied to a chromosome that has its centromere in the middle

MetadataThe details, usually in digital form, of experimental conditions and the procedures followed for the phenomics studies (or any other study).

Metaphase. The stage of mitosis or meiosis at which chromosomes align along theequatorial plane of the cell.

Metaphase: A stage in mitosis or meiosis during which the chromosomes arealigned along the equatorial plane of the cell.

Meta-QTLsQTLs identified by QTL meta-analysis. Also called true QTLs.

Metaxenia. The influence of pollen on the maternal tissues of the fruit. It greatlyinfluences fruit quality in date palm.

Methylation. Modification of a molecule by the addition of a methyl group. Many amutagen acts through methylation (e.g., MMS), causing mispairing between bases,and eventually leading to GC 􀄺 AT transition.

Metroglyph Analysis (Anderson 1957). A semi-graphic method of studying variabilityin a large number of germplasm lines taken at a time. For variability assessment, thefirst two characters, which are highly variable, are depicted on X and Y-axes,separately. The mean values of X (say, character-1) for each genotype is plottedagainst the mean values of Y; thus, each line occupies a definite position on the graph,called glyph. Variation for the remaining characters of each genotype is displayed onthe respective glyph by distinctive rays. Each character occupies a definite rayposition. The length of the ray for a particular character on the glyph may be short,medium, or long depending on the index value of a genotype. It is a very easytechnique to study the pattern of morphological variation among genotypes, and canbe applied to both replicated and unreplicated data. However, inclusion of a largenumber of genotypes sometimes leads to overlapping of glyphs on the graph.

Microarray: A large set of cloned DNA molecules spotted on to a solid matrix (such as a microscope slide) for use inprobing a biological sample to determine the gene expression, marker pattern, or nucleotide sequence of DNA/RNA.

MicroarrayA small plaque/wafer of silicon, glass, or metal, onto which one end of multiple copies of each of a large number of different single-stranded DNA molecules is covalently linked and the different molecules are arranged in separate spots.

Microarrays: A tool to examine the expression levels and intertwinedinteractions among genes and their products; complementary DNA of interest isaffixed to a glass slide in an ordered array; the expression level is determinedby the binding to cDNA; a small glass, or filter, square may contain probes forthousands of gene products; in statistics, a microarray is a block design with blocksof size 2; the concept arose in DNA testing, where each array is probed with twoDNA samples

Microcentres (Harlan 1951). The small regions enriched with tremendous plantdiversity and rapid rate of plant evolution. These microcentres may offer excellentopportunities to collect valuable types, and to study evolution of cultivated plantsexperimentally. (Also see ecological regions).

Micro-mutation. A mutation with a small effect that can be detected only bymeasurement on a group of plants.

Micron C7. The smallest known gene, also designated by Mcc C7. It produces aheptapeptide inhibiting protein synthesis in the Enterobacteriaceae. The heptapeptideis synthesized from 21 bp open reading frame.

Micronaire. An instrument that measures the rate of airflow through a standard volumeof cotton. From its readings, fibre fineness and maturity of cotton samples may becompared.

Micropropagation. In vitro propagation of plants from very small tissues (parts). It isone of the best and most successful examples of the commercial application of tissueculture technology. Advantages of micropropagation include: (a) identical progenytrue-to-the mother plant, (b) rapid and large-scale multiplication of elite genotypes,and (c) rejuvenation of old clones/varieties. Besides, it is season-neutral andfacilitates germplasm exchange by avoiding the risk of spreading pathogens andinsect-pests.

Micropropagation: In vitro clonal propagation; the multiplication of geneticallyidentical copies of a cultivar by asexual means is called clonal propagation

Micropyle. The usual point of entry for pollen tube in the embryo sac.

Microsatellite marker: Microsatellites or simple sequence repeats are a typeof molecular marker; microsatellites consist of tandem repeats of 1–6 nucleotidemotifs; the repeats usually are in units of ten or more, although repeats as small assix units have been found

Microsatellite. A type of repetitive DNA based on very short repeats such asdinucleotides.

Microsatellite: A repeated motif of nucleotides, usually onlytwo or three bases in length, where the number of repeatsfrequently differs between different members of a species.

Microsatellite-primed PCR (MP-PCR): A technique resulting in RAPD-like patterns after agarose gel electrophoresis and ethidium bromide staining; theMP-PCR technique is more reproducible than RAPD analysis because of higherstringency

MicrosatellitesUsually, <100 bp long sequences comprising tandem repeats of 2–7 bp.

Microspecies. A small breeding population with limited variability. It may be a biotypeof the species.

Microspore Mother Cell. The diploid cell (2n) in the anther, which gives rise, throughmeiosis, to four haploid microspores.

Microspore. One of the four haploid spores originating from meiotic division ofmicrospore mother cell in the anther and which gives rise to the pollen grain.

Microsporocyte. The microspore mother cell (or pollen mother cell).

MicrosyntenySynteny based on DNA sequence.

Midparent Value. The mean value of a quantitative phenotype obtained from twospecific parents; the arithmetic mean of two parents with respect to a trait(s). Forinstance, if a parent is 20 unit on the phenotypic scale of measurement and the secondone 30 unit, the midparent value will be 25 unit [MP=(20+30)/2].

Migration. Any form of introduction of genes (individuals) from one population intoanother. It can be responsible for introducing new genes or moving up or downwardsthe frequency of genes already present in the breeders’ or natural populations. Thechange in frequency of the gene already present is expressed as: 􀇻p = m (P-pt),whereas, m is the proportion of migrants, P is frequency in donor population, and pt isfrequency in the recipient population. Clearly there will be no change in the gene frequency, if there is no difference in gene frequency between donor and recipientpopulation.

Mini Core. See core collection.

Minichromosome: A very small chromosome, usually as a result ofchromosome aberrations; engineered minichromosomes offer an opportunity toimprove crop performance; unlike conventional gene transformation technologies,minichromosomes can be used simultaneously to transfer and to stablyexpress (multiple) sets of genes; because they segregate independently of hostchromosomes, they provide a platform for accelerating plant breeding; strategiesfor producing artificial chromosomes may consider engineering of endogenouschromosomes or de novo assembly from chromosomal constituents

Minimal Medium. A medium containing only a carbon source (sugar), inorganic saltsand water.

Minisatellite. A type of repetitive DNA sequence based on short repeat sequences with aunique common core. It is used for DNA fingerprinting.

Minisatellite: Highly polymorphic DNA markers comprised of a variablenumber of tandem repeats that tend to cluster near the telomeric ends ofchromosomes; the repeats often contain a repeat of 10 nucleotides; they are usefultools for genetic mapping

MinisatellitesSequences, typically 0.2–2 kb long, made up of 11–60 bp long tandem repeat units having identical or almost identical sequences.

Minor QTLA main effect QTL that explains less than 10 % of the phenotypic variance for the concerned trait.

Mispairing. The presence of a non-complementary nucleotide at a given position in oneof the polynucleotide chains of a DNA double helix. It may lead to a gene mutation.

Missense Mutation. A mutation that alters a codon in such away that it encodes adifferent amino acid, for example, GAA (glutamic acid) to GUA (valine). Thischange, which occurs in the B polypeptide chain at 6th amino acid position, causessickle cell anaemia in human population.

Missense mutation: A single DNA base change which leads to a codonspecifying a different amino acid.

Missing heritabilityThe part of phenotypic variation in a quantitative trait that is not explained by the QTLs identified by various AM studies.

Missing Plot Technique. A technique to calculate the value of any missing treatment inany block (replication). The calculation for any missing value (x) is done as: x = [r Bj+ t Ti – GT / (r-1) (t-1)]; whereas, r = no. of replication, t = no. of treatments, Bj =known jth block total where x is missing, Ti = known treatment total for which x ismissing, and GT = grand total with missing observation. Missing plot technique is anadvantage for a randomised complete block design.

Mitochondrion. A eukaryotic cell organelle that is the site of ATP synthesis, and of thecitric acid cycle. It is a small body usually with distinct shelf like internal layers. Itprovides energy through respiration and oxidation.

Mitogen. Any substance that stimulates cells to undergo mitosis.

Mitosis. A kind of cell division in which the nucleus is divided into two daughter nucleiwith equivalent chromosome complements. It is a conservative process that maintainsthe same chromosome number in the daughter cells compared to the parental one. Itoccurs in both somatic as well as reproductive cells.

Mitosis: The process of nuclear division in cells which producesdaughter cells that are genetically identical to eachother and to the parent cell.

Mitotic Crossover (Stern 1943). A crossover resulting from the pairing of homologsduring mitosis; also called somatic crossover. Like meiosis, it takes place in fourstrandstage of chromosomes. The outcomes of somatic crossing-over depend upon:147(a) the site of the crossing-over, and (b) the mode of orientation and, thus distributionof the centromeres of the chromosomes engaged in somatic crossing-over. It alwaysoccurs in the diploid cells. It can be an important source of variation in asexualpathogenic fungi. It is also thought to be important in allowing recessive cancercausingmutations to become expressed.

Mitotic recombination: The recombination of genetic material during mitosisand the process of asexual reproduction; the mechanism for the production ofvariation in heterokaryons

Mixed linear modelThe markers and the population structure (Q) are treated as fixed linear effects and the additive effects of the multiple background QTLs are considered as linear random effects.

Mixoploidy. A term that covers all types of chimeras in which the heterogeneity betweendifferent elements involves differences in chromosome number (2n = x, 2x, 3x, 4x, 2x-1, 2x-2, etc)

Mode. The value of the variate, which occurs most frequently in a data set. In a frequencytable, the modal class is the class that has the greatest frequency.

Model organisms: Creatures used in genomic analysis because they have many genetic mechanisms in common with each other and with humans. Theseorganisms lend themselves well to classical breeding experiments and directmanipulation of the genome.

Model-based clustering methodsCluster membership is based on some parametric evolutionary model.

Modified Convergent Cross (Mackey, 1954). A convergent cross that involves a verygood variety as persistent crossing partner at every stage of the crossing procedure.This results in 50% of the genes from the persistent good variety. For example:A × B A × C A × D A × E F1 × F1 F1 × F1 F1 × F1F1(50% genes from the parent A)The procedure can be combined further with different degrees of back crossing(convergent back cross) of F1’s with the desired parent (A) to ensure even higherpercentage of genes from the very good variety (A).

Modified Doak Method (Mehta and Patel 1983). A method of producing hybrid seed incotton without emasculation. The top portion of the corolla is impressed deeply toexpose only the tip of the stigma. The anthesis is prevented through placing lintdipped in mud over the remaining portion of the corolla. The stigma is pollinated inthe next morning. The boll setting in crosses has been found greater (specially in thediploid cotton) than with Doak Method.

Modified Single Cross. The progeny of a cross between a single cross, derived from tworelated inbred lines, and an unrelated inbred line, (A1 × A2) × B.

Modifier. A gene(s) that affects the expression of other gene(s) primarily in a quantitativemanner. It can act as enhancer or inhibitor depending upon the situation. Its effectmay be dominant or recessive. The degree of dominance or recessiveness of a genemay also be influenced by modifying genes. Modifiers may have a large or smalleffect (but the mode of action is almost always quantitative). Some of them may havetheir own primary function and modifying actions appear as secondary effects, whileothers have solely enhancing or inhibitory effects on the expression of other genes.The well-documented example is that of spotting in mice. Similar cases in plants doexist. Actually further improvement in any character (wherein major genes havealready been exploited) will depend upon manipulation of modifiers.

Molecular beaconsSpecially designed oligonucleotide hybridization probes used for identification of SNP alleles.

Molecular Biology. A branch of modern biology in which biological phenomena arestudied by physical, chemical and biochemical investigations at the molecular level.

Molecular biology: The study of the structure and function of proteins andnucleic acids in biological systems.

Molecular biomarkersThose dynamically expressed molecules that can be measured and used as indicators of specific phenotypic features.

Molecular breeding: Plant breeding assisted by using DNA markers or proteinmarkers.

Molecular Cytogenetics. A correlated study of cytology and genetics at molecular level.It may provide clues about the stability and expression of trans genes.

Molecular farming: The application of biotechnology for the selected productionof pharmaceutical compounds or other health or industrial compounds within aliving organism (eg. microbe or agricultural crop)

Molecular Genetics. The study of genetic systems that can be described at the molecularlevel. It is, therefore, the study of molecular processes underlying gene structure andfunction.

Molecular genetics: The study of genes, the base components of heredity. Genesare made up of DNA, whose bases are placed in a unique order that determinesthe hereditary traits in living organisms.

Molecular inversion probeA single 120 nucleotide (nt) long oligonucleotide that hybridizes to a specific sequence of the genome and forms a circle that has a single base pair gap at the SNP site. The assay involves primer extension and ligation producing a closed circular molecule.

Molecular marker: A gene or DNA sequence that identifies a particular locuson the chromosome (whether the actual location is known or not) and whose inheritance can be followed. An identifiable physical location on achromosome (e.g., restriction enzyme cutting site, gene) whose inheritance can be monitored.

Molecular Markers. Agents that mark genetic variation at DNA level. There arenumerous molecular markers, for example, RFLP, RAPDs, AFLP, STS, EST, SSRs(also called microsatellites), SCARs, and the like. Except RFLP (that is southernhybridisation based molecular marker), all are PCR-based markers. AFLP combinesthe properties of RFLP (restriction fragment length polymorphism) and PCR(polymerase chain reaction). However, STS (sequence tagged sites) is more desirablebecause of high degree of reproducibility and large-scale automation, which isessential for handling a large number of samples. These are numerous, permanent anduninfluenced by environments/developmental stage of plants. Molecular markers canbe reliably used in the selection of superior genotypes. However, these are expensivecompared to morphological or biochemical ones. These can be used to: (a) develop149saturated maps, (b) fingerprint DNA for varietal characterization, (c) studyphylogenetic and evolutionary relationship among related species, (d) characterize A,B, and R lines and find mechanisms of heterosis, (e) tag gene/ genes of interest, (f)practice marker-assisted selection, and (g) map orthologous gene(s). Markers may beof two types: screenable and selectable. The former allows screening of transformedcells through expression of specific enzyme to produce phenotype enabling us toidentify transformed cells. But selectable markers are genes that confer resistance tosome compounds like herbicides and antibiotics usually toxic to normal plants; thusonly cells having these markers survive under selective conditions. It is emphasizedthat the importance of marker loci is solely that they identify “short chromosomesegments” that carry predominantly favourable alleles. However, such chromosomesegments must be small so that it may remain intact through many cycles of crossingover and segregation. This is true for marker loci marking even QTLs.

Monad. An individual cell produced by a meiocyte (instead of tetrad) as a result ofmeiotic abnormality.

Monocentric chromosome: A chromosome with only one centromere

Monocistronic mRNA. An rnRNA that encodes only one protein.

Monoclinous. Having male and female germ cells in one and the same flower.

Monoecy. A condition/system wherein staminate (male) and pistillate (female) flowersare borne separately on the same plant (for example, corn).

Monogenomic Species. Species which are basically diploid containing only a single kindof genome. B. campestris (AA), B. nigra (BB), and B. oleracea (CC) are some of theexamples. Such species are also called primary species.

Monogeny. The production of only male or female offspring.

Monogerm. A sugar beet seed with a single germ, in contrast to a multigerm seed.

Monohaploid: A haploid cell or individual possessing only one chromosomeset in the nucleus

Monohybrid Cross. A cross between two individuals identically heterozygous at onelocus, Aa × Aa, for example.

Monohybrid. A hybrid that is heterozygous with respect to one gene only, Aa, forexample. The hybrid at two loci (AaBb) is called a dihybrid, and so on.

Monomer. A polypeptide.

Monophyletic Evolution. Evolution of individuals from a single interbreedingpopulation.

Monoploid. An organism having a single complement of a basic chromosome set of thespecies; an individual having a single genome of the species. For instance, monoploidof bread wheat may have either A, B, or D genome. It (monoploid) may arisespontaneously in nature. The surviving monoploids look lean and thin. Sincechromosomes do not have pairing-partners, random segregation occurs duringmeiosis. Gametes are nearly always deficient in one or more chromosomes, and thusmonoploids are usually sterile. For instance, a monoploid of barley has sevenchromosomes. A functional gamete may be formed if it has all the sevenchromosomes. However, the probability of going all the seven chromosomes to one ofthe poles during meiosis I is (½)x-1 = (½)7-1 = (½)6. This amounts to the probabilityof a functional gamete. Thus the probability that a monoploid of barley sets one seedis (½)6× (½)6 = (½)12. This explains why a monoploid plant is almost sterile.

Monosome. A chromosome with no homolog to pair with; a single chromosome in anotherwise diploid individual (2n-1).

Monosomic. An individual lacking in one chromosome of the diploid complement,henece, having 2n-1 chromosomes. The missing chromosome is called a monosome.In monosomics, the unpaired chromosome passes at random to either pole duringmeiosis; however, it frequently lags at anaphase, and is not included in either daughternucleus. For this reason, gametes with n-1 chromosomes are frequent compared tothose with n chromosomes. However, this bias is not reflected strikingly in thezygotic chromosome numbers because gametes with n-1 chromosomes often do notfunction (especially 􀆃 gametes). Furthermore, zygotes with 2n-2 chromosomes areinviable except in a few polyploid species. Thus most of the progeny of monosomicsare either normal diploids (2n) or monosomics (2n-1).

Monosomic: A genome that is basically diploid but that has only one copyof one particular chromosome type, so that its chromosome number is 2n–1; monosomic series were developed

Monosomy: The total loss of one of a pair of chromosomes. This occurs, forexample, in Turner Syndrome where one X chromosome is lost leaving a total of45 chromosomes.

Monotelocentric: A cell or individual lacking one chromosome pair butshowing one telocentric chromosome for one arm of the two missing homologues

Monotelodisomic: A cell or individual lacking one chromosome pair butshowing two homologous telocentric chromosomes for one arm of the two missinghomologues

Monotelomonoisosomic: A cell or individual lacking one chromosome pairbut showing a telocentric chromosome for one arm of the missing homologouspair and an isochromosome for

Monotelotrisomic: A cell or individual showing an additional telocentricchromosome to a certain pair of chromosomes more abundant parents but alsopossess some of the characters of the other parent species

Morgan: A measure of genetic distance; one Morgan (M) equals 100 centimorgans.

Morph. Any one of the genetic forms that account for polymorphism.

Morphogenesis. A developmental process that leads to changes in gross form, cellularfine structure or both; processes giving final shape to an adult organism byestablishment of specific pattern of tissues and organs.151Mosaic. Also called a chimera (more frequently in plant species). A tissue containing twoor more genetically distinct cell types or an individual composed of such tissues.

Morphological markers: Simply inherited and easily scored morphological traits; the earliest genetic markers.

Mosaicism: Where a genetic or chromosomal abnormality does not occur in allbody cells. Often related to X chromosome innactivation.

Moving Means Method (Knott 1972, Townley-Smith and Hurd 1973). A method oftesting the genetic potential of early generation lines (F2/F3/F4) in unreplicated plots.It involves taking mean of a number of adjacent plots (excluding the plot in question)as an environmental index. The performance of a test genotype is expressed as thedifference between its own value and the moving mean, or as the percentage ofmoving mean.12.00 13.00 15.0018.00 20.00 17.0014.00 19.00 15.00In the above example, the value of moving mean is{(12+13+15+18+17+14+19+15)/8} = 15.38. Thus, performance of the test genotype(middle plot) = (20.00/15.38) × 100 􀂧 130%. Its control over experimental error iscomparable to the use of intermittent check plots; however, it has an added advantagesince no plots are wasted for accommodating the check entries.

mRNA. An RNA molecule transcribed from the DNA of a gene under the influence of anenzyme transcriptase. The genetic message is translated into a polypeptide by theaction of ribosomes. During the process, only the genetic message flows from DNA toRNA (there is no physical transfer of any material).

Multiline (Jensen 1952). A blend of compatible lines, each selected for similarity ofheight, maturity, and other agronomic or horticultural characteristics, but carrying adifferent gene(s) for resistance. However, Norman Borlaug (1954) has suggested asimilar but somewhat more sophisticated approach based on lines developed bystandard back cross breeding. In the present context, a multi-line cultivar is a mixtureof isogenic lines. In this approach, genes for specific resistance (that reduce the initialamount of inoculum) are considered in the component lines. However, in theaftermath, the overall result appears as horizontal resistance, since the rate ofpathogen increase is also considerably reduced. Browning and Frey (1969) have152called it synthetic horizontal resistance or population resistance. It is only a concept;now-a-days, it is not used practically.

Multiline Variety. A variety developed through a composite of isolines. The term“multilines” sometimes is applied to mixtures of genetically diverse lines produced invarious ways to buffer against environmental stresses. More accurately, thesepopulations should be called composites instead. Such a variety is now rare incultivation.

Multiline: A mixture of isolines, each of which is different fora single gene conditioning different forms of the same trait.

Multilocus mixed model: It includes multiple loci as cofactors in the AM model and employs a simple stepwise mixed-model regression analysis combined with forward inclusion and backward elimination of loci in the model.

Multimer. A protein consisting of two or more subunits (monomers or polypeptides).

Multimeric Structure. A structure composed of several identical or different subunitsheld together by weak bonds.

Multiparent advanced generation intercross populations: A collection of RILs produced from a complex crossbred/outbred population involving several parental lines.

Multiple Allele. A member of a series of several known allelic forms of a gene. Theexistence of such a situation is called multiple allelism. The presence of multiplealleles has been observed for the self-incompatibility system in the Nicotiana spp.Please notice that the presence of all the allelic forms of a gene in any naturalpopulation is a rare event. As the no. of allelic forms increase, the possible kinds ofgametes (and homozygotes) and total no. of genotypes increase exponentially. Forexample, for a single gene pair with four alleles, the kinds of gametes (homozygotes)and total no. of genotypes will be 41 (= 4) and (4 ×5)/2 (= 10), respectively. For twogenes with same no. of alleles at the two loci, the values will be 42 (= 16) and(16×17)/2 (=136), respectively.

Multiple Correlation Coefficient. A statistic measuring the joint association of all theindependent variables with the dependent one. It thus tells how much of the variationin the dependent (Y) could be accounted for by reference to these independent ones.Numerically, it is the square root of the ratio of the regression sum of squares to thetotal sum of squares, and is denoted by R.

Multiple Cross. A cross involving many parents. Theoretically its objective is to obtainrecombination from many a parent; the practical utility of multiple cross, however, islimited in plant breeding for two reasons: (a) the size of the population increasesexponentially with increasing no. of parents to ensure occurrence of the desiredgenotype in the segregating generation, and (b) it is often difficult to find many welladapted parents having different useful genes. (Also called convergent cross).

Multiple interval mapping: An approach for simultaneous QTL mapping in multiple marker intervals.

Multiple QTL mapping: It combines simple interval mapping with multiple regression analysis and includes all the significant QTLs in the genetic model used for mapping.

Multiple Resistance. A strategy in resistance breeding that involves placement of two,three, or more new and still effective resistance genes into a new cultivar to impose154barriers of several resistance to the pathogen population simultaneously. It should bean effective strategy because a new race, to overcome multiple resistance genes, musthave two or more simultaneous changes towards the virulence, whereas a new raceneeds only one change towards virulence to overcome a single gene for resistance. Itcan be effective if the breeding is coordinated centrally, and the production area isisolated from other areas where the system is not applied.

Multiple-Factor Hypothesis (Nilsson-Ehle 1908). A hypothesis to explain quantitativevariation. Many genes, each with a small but equal effect, segregate to produce acontinuous variation in a quantitative character. Conclusive evidences for thishypothesis proved beyond doubt that all genetic traits (whether qualitative orquantitative) follow Mendel’s laws of heredity.

Multiplex PCR: Two or more primer pairs used for amplification of two or more loci in a single PCR reaction tube.

Multiplex ratio: The average number of markers scored per assay of a marker system in a given population.

Multiplexed shotgun genotypingSize-selected (250–300 bp) restriction fragments from several individuals are separately ligated to distinct barcodes, pooled, and sequenced using an NGS platform.

Multiplex-endonuclease genotyping approach AFLPA modification of AFLP; four or more endonucleases are used for the digestion of the sample DNA; only one pair of adapters used for amplification.

Multiplexing: A sequencing approach that uses several pooled samplessimultaneously, greatly increasing sequencing speed.

MultiplexingCarrying out two or more different reactions, e.g., PCR amplification, in a single tube or separating the products of two or more PCR reactions in a single gel lane.

Multispectral reflectance data Reflectance data acquired at few selected wavelengths.

Multitrait introgressionIntrogression of genes governing two or more different traits into a single RP.

Multitrait mixed model: It extends the linear mixed-model approach of AM to the analysis of pairs of correlated traits.

Multivalent. A pairing configuration of three or more completely or partiallyhomologous chromosomes observed during meiosis-I. Multivalent formation is acharacteristic feature of autopolyploids.

Multivariate linear mixed modelIt allows testing of associations between markers and multiple correlated phenotypes and is able to control population structure.

Multivariate methodsMethods for the analysis of data on multiple traits for each entity.

Mutagen Effectiveness. The number of induced mutations per unit dose of a mutagen.

Mutagen Efficiency. A ratio of specific desirable mutagenic change(s) to the undesiredeffects such as plant damage, sterility, lethality, and the like.

Mutagen. An agent (physical, chemical or even biological) that is capable of increasingthe mutation rate.

Mutagen: Any agent that is capable of inducing and increasing the mutation rate.

Mutagenesis. Treatment of plants or plant parts with a mutagen to increase mutationrates.

Mutant Allele. An allele differing from the allele found in the standard or wild type.

Mutant allele: An allele differing from the allele found in the standard, or wildtype.

Mutant Hunt. The process of collecting different mutants showing abnormalities in acertain structure or in a certain function, as a preparation for mutational dissection ofthat function.

Mutant Site. The damaged or altered area within a mutated gene.

Mutant. An organism or cell carrying a detectable mutation. The term is an adjective;thus it must precede a noun (a mutant individual, for example).

Mutant: A gene having undergone genetic change or mutation.

Mutation Breeding. A system of breeding in which plants/plant parts/seeds or any kindsof propagules are treated with any kind of mutagen accompanied by selection fordesired types in succeeding generations.

Mutation Event. The actual occurrence of a mutation in time and space.

Mutation Frequency. The proportion of mutants in a population.

Mutation Pressure. The continued recurrent mutation tending to increase the frequencyof the mutated allele in the gene pool of a population.

Mutation Rate. The number of mutation events per gene copy in a population per unit oftime (for instance, per cell generation).

Mutation. In broadest sense, any heritable variation in a gene or in chromosome structureand/or number. However, the use of the term has now been restricted to include onlygene mutation, which refers to a process by which new allele(s) is produced throughheritable structural changes in the gene(s). It may be the process or even the result ofthe process. It may occur in nature spontaneously, but the frequency is very low(about one per million). The rate of mutation can be increased in breeders’populations through chemical or physical agents called mutagens. The molecularbasis of mutation relates to mispairing between nucleotides of the DNA moleculeirrespective of whether the mutation is natural or artificial. In populations of a species,it the ultimate source of all variation. If an allele A mutates to a (ignoring backmutation) with the mutation rate 􀈝 for a given number of generations n, thenfrequency of the allele A after nth generation, pn = p0e-n􀈝 (whereas, e is base of thenatural logarithms). Thus the frequency of the A allele decreases with time but therate will be very slow. Therefore, the process of mutation cannot drive the process ofevolution unless supplemented with recombination or migration. The statement that amutation, if not favoured, is lost from a population is not true absolutely. Theprobability that it could be lost is (2N-1)/2N (N = size of the population); if not, thenthe probability that it is fixed is 1/2N. Thus a mutation can become established in apopulation even though it is not favoured by natural selection simply by a process ofrandom genetic drift.

Mutation: A permanent change in the genetic material involving either a physical alteration in the chromosome or a biochemical change in the underlying DNA molecule.

Mutational Dissection. The study of the components of a biological function through astudy of mutations affecting that function.

Mutational Equilibrium. The product of frequency of mutations per individual pergeneration and average number of generations during which mutated gene ismaintained in the population before it is eliminated.

Mutator Gene. Any gene that increases mutation rate of other genes.156

Mutein. A protein with one or more mutationally altered amino acids in thepolypeptide(s). it is analogous to normal or wild type protein. It may or may not lackenzymological, immunological, or physio-chemical activity.

MutMap schemeA quick, reliable, and cost-effective method for mapping of causal SNPs in induced mutations. It uses a single bulk of the mutant plants from the F2 generation of mutant parent cross and a reference genome for alignment of the NGS sequence data from this bulk.

MutMap-Gap scheme It identifies causal mutations located in the genomic regions missing from the parental/reference genome.

Muton. The smallest part of a gene that is involved in a mutation event. It is now knownto be a nucleotide pair.

Mycoplasma. Small bacteria that produce infectitious diseases in plants and animals.Among living organisms, they have the smallest mass. Mycoplasma can be cultured invitro like any bacteria. Mycoplasma genitalium with a genome size of 580 kb DNA isthe first complete life.


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