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Inversions result when there are two breaks in a chromosome and the detached segment becomes reinserted in the

reversed order. They are classified into two types depending upon the inclusion or absence of the centromere within the inverted segment. Thus when both breaks occur in one arm of the chromosome it leads to a paracentric inversion; when a break occurs in each of the two arms, the centromere is included in the detached segment and leads to a pericentric inversion. Meiosis is normal in inversion homozygotes. In heterozygotes pairing between homologous chromosomes is affected in the region of the inverted segment. Consequently, there is a suppression of recombination and fertility is impaired. Paracentric inversions: This type of inversion is identified in the heterozygote by formation of a pairing loop at pachytene. If the size of the loop is large enough, chiasma formation will take place within it. When a single chiasma forms between an inverted and a normal segment, the two chromatids involved will produce one dicentric chromatid and one acentric fragment after exchange (Fig. 12.11). The other two chromosomes will be normal. At anaphase I the dicentric chromosome will be pulled towards both poles, it will form a bridge that will ultimately break. The acentric fragment due to its inability to move would be eventually lost. Consequently, of the resulting four gametes, two would be normal and two deficient in chromosome segments. In plants deficient gametes are not viable (pollen grains that are deficient usually abort and are nonfunctional). In animals such gametes take part in fertilisation but either the zygote or the embryo aborts. In an individual heterozygous for a paracentric inversion therefore, viable offspring are produced only by two of the four chromatids which did not have chiasma formation between them in the region of the loop. In each chromatid the gene sequence in the inversion segment will be of the nonrecombinant, parental type. Consequently, none of the offspring would be recombinants for genes present within the inverted segment. In this way a paracentric inversion suppresses recombination throughout its length. In some insects and in Drosophila, individuals heterozygous for an inversion do not

show reduction in fertility. In fact paracentric inversions occur frequently in natural populations of Drosophila. There are two explanations for this. One is absence of crossing over in male meiosis. The second is occurrence of four products of female meiosis in linear order of which the middle two egg nuclei have the deficiency; the peripheral two nuclei are functional and fertilised. They produce viable offspring of the parental type.

Inversions with such devastating effects can be traced to meiosis, when a crossover occurs between the inverted chromosome segment and the noninverted homolog. To allow the genes to align, the inverted chromosome forms a loop. When crossovers occur within the loop, some areas are duplicated and some deleted in the resulting recombinant chromosomes. In inversions, the abnormal chromosomes result from the chromatids that crossed over. Two types of inversions are distinguished by the position of the centromere relative to the inverted section. A paracentric inversion does not include the centromere ( figure 13.20 ). A single crossover within the inverted segment gives rise to two normal and two very abnormal chromosomes. The other two chromosomes are normal. One abnormal chromosome retains both centromeres and is termed dicentric. When the cell divides, the two centromeres are pulled to opposite sides of the cell, and the chromosome breaks, leaving pieces with extra or missing segments.

The second type of abnormal chromosome resulting from a crossover within an inversion loop is a small piecethat lacks a centromere, called an acentric fragment. When the cell divides, the fragment is lost because acentromere is required for cell division.

Pericentric inversions: In an individual heterozygous for a pericentric inversion, the centromere is present within the loop. When chiasma formation takes place within the inverted segment the chromatids resulting after exchange do not form a dicentric and acentric fragment as in a paracentric inversion heterozygote. Instead, they have one centromere each, but are deficient for some segments whereas other segments are duplicated (Fig. 12.12). The exchange segments produce inviable gametes and offspring. As in the case of pericentric inversions, the two chromatids not involved in crossing over only produce viable offspring with parental combination of genes present in the inverted segment.