CHROMOSOMES

E straussberger (1875) discovered thread like structures during cell division and called chromosomes (chroma=colour) due to their affinity to basic dye. In eukaryotes, well organised nucleus contain definite number of chromosomes of definite size and shape. At leptotene stage of meiotic prophase chromosomes appear as headed structure called chromomeres, size of chromomeres and interchromomeric regions are not constant, so that every leptotene has its own particular pattern. The DNA is though known to concentrate in the chromomeres, but it is believed to be present in interchromomeric region also. Number, size and shape of chromosome: Chromosome number varies from 2n = 4 in Haplopappus gracilis (compositae) to 2n = >1200 in some pteridophytes, Aulacantha, a radiolarian, 2n = 1600 n signifies gametic or haploid chromosome number 2n signifies somatic or diploid chromosome number In polyploids basic chromosome number (x) Chromosome is normally measured in mitotic metaphase as short as 0.25 in fungi and birds as long as 30 in some plants like Trillium, Drosophila 3, Humans 5, Maize 8-12 . Chromosome shape is usually observed at anaphase, when the position of centromere/ primary constriction determines chromosome shape. Centromere can be terminal, sub terminal or median

Terminal Sub-terminal Median Sub median Morphology :

Rod shape J shape V shape L shape

Corresponding to different position of centromere Acrocentric or Telocentric Submetacentric Metacentric Secondary constriction can be observed in some chromosomes and forms a satellite. It remains attached to rest of the body by a thread of chromatin. Secondary constrictions are constant on their position and can be used as useful markers. They can be distinguished from centromere because chromosome bends or angular deviation only at the position of centromere, chromosomes having satellite are called as SAT-Chromosomes. The chromosome extremities or terminal regions on either side are called telomeres. Detailed study of chromosome morphology reveals a coiled filament throughout the length of the chromosome. This filament is called chromonema (Vejdorslay, 1912). The chromonemata form the gene-bearing portions of the chromosomes. The chromonemata are embedded in the achromatic substance known as matrix. Matrix is enclosed in a sheath or pellicle. Both matrix and pellicle are non-genetic materials and appear only at metaphase when the nucleolus disappears.

 It is believed that nucleolar material and matrix are interchangeable i.e. when matrix disappears, nucleolus appears and vice versa. Chromatid is a half chromosome & two chromatids being connected at centromere. The chromonema is a sub-chromatid structure and there can be more than one chromonemata at the centromere.

Karyotype :
Group of characteristics like chromosome number, size and shape of a chromosome set up of a species is called as karyotype and is usually represented by a diagram called as idiogram, where chromosomes of haploid set are ordered in a series of decreasing size. Karyotype are presumed to represent evolutionary relationships. Karyotype suggests primitive or advanced features of an organism. Asymmetric karyotype - karyotype showing large difference between smallest and largest chromosome of the set and having fewer metacentric chromosomes. It is considered to be a relatively advanced features when compared with symmetric karyotype. Levitzky (1931) suggested that in flowering plants there is a predominant trend towards karyotype asymmetry. When chromosomes stained with acetocarmine or feulgen (basic fuchin), the darkly stained heterochromatic and lightly stained euchromatic regions can be differentiated. Heterochromatin is found at chromomeres, chromocenters and knobs. Chromomeres are regular feature of all prophase chromosomes, but their number, size, distribution and arrangement are specific for a particular species at a particular stage of development.

Human male karyotype

Human karyotype

 Chromocenters are heterochromatic regions of varying size which occurs near the centromeres in proximal regions of chromosome arms. Knobs are spherical heterochromatin bodies which may have a diameter equal to there chromosome width but may reach the size having diameter which is several times the width of chromosomes. Very distinct chromosome knobs can be observed in maize at pachytene stage. Knobs are valuable chromosome markers for distinguishing chromosomes of related species and races. Constitutive heterochromatin - certain regions of chromosomes particularly those proximal to centromeres are constant and are called constitutive chromatin. Facultative heterochromatin - other heterochromatin regions and represented by whole sex chromosome which become heterochromatic only at certain stage. Eg. female humans - one X chromosome is inactivated facultatively. Plant - accessory chromosomes are heterochromatic. Dioecious genera - Melandrium and Rumex - one or both sex chromosomes may undergo partial or complete heterochromatization. Y chromosome - heterochromatic having inactive genes in several dioecious plants and animals. Nucleosome model - R.D Kornberg and J.O Thomas (1974) term nucleosome - P. Oudet (1975) Folded fibre model - E.J dupraw (1965)

Chromomeres in polytene chromosomes

Chromocenter

The BRM protein is associated with chromosome puffs and interbands of salivary gland polytene chromosomes. (A) Distribution of BRM protein on wild-type polytene chromosomes. Arrowhead indicates chromocenter. (B) The top panel shows indirect immunofluorescence using an anti-BRM antibody (green), the second panel is DAPI-stained DNA (blue), the third panel shows the merged images and the bottom panel is a split image. Note that BRM protein is predominantly found in the DAPI interbands. The distal region of chromosome arm 2L is shown.