THE EVOLUTION OF HETEROMORPHIC SEX CHROMOSOMES

The facts and ideas which have been discussed lead to the following synthesis and model. 1. Heteromorphic sex chromosomes evolved from a pair of homomorphic chromosomes which had an allelic difference at the sex-determining locus. 2. The first step in the evolution of sex-chromosome heteromorphism involved either a conformational or a structural difference between the homologues. A structural difference could have arisen through a rearrangement such as an inversion or a translocation. A conformational difference could have occurred if the sex-determining locus was located in a chromosomal domain which behaved as a single control unit and involved a substantial segment of the chromosome. It is assumed that any conformational difference present in somatic cells would have been maintained in meiotic prophase. 3. Lack of conformational or structural homology between the sex chromosomes led to meiotic pairing failure. Since pairing failure reduced fertility, mechanisms preventing it had a selective advantage. Meiotic inactivation (heterochromatinization) of the differential region of the X chromosome in species with heterogametic males and euchromatinization of the W in species with heterogametic females are such mechanisms, and through them the pairing problems are avoided. 4. Structural and conformational differences between the sex chromosomes in the heterogametic sex reduced recombination. In heterogametic males recombination was reduced still further by the heterochromatinization of the X chromosome, which evolved in response to selection against meiotic pairing failure. 5. Suppression of recombination resulted in an increase in the mutation rate and an increased rate of fixation of deleterious mutations in the recombination-free chromosome regions. Functional degeneration of the genetically isolated regions of the Y and W was the result. In XY males this often led to further meiotic inactivation of the differential region of the X chromosome, and in this way an evolutionary positive-feedback loop may have been established. 6. Structural degeneration (loss of material) followed functional degeneration of Y or W chromosomes either because the functionally degenerate genes had deleterious effects which made their loss a selective advantage, or because shorter chromosomes were selectively neutral and became fixed by chance. 7. The evolutionary routes to sex-chromosome heteromorphism in groups with female heterogamety are more limited than in those with male heterogamety. Oocytes are usually large and long-lived, and are likely to need the products of X- or Z-linked genes. Meiotic inactivation of these chromosomes is therefore unlikely. In the oocytes of ZW females, meiotic pairing failure is avoided through euchromatinization of the W rather than heterochromatinization of the Z chromosome.(ABSTRACT TRUNCATED AT 400 WORDS)     

THE CHEMICAL COMPOSITION OF ISOLATED CHROMOSOMES

By means of 1 M NaCl isolated lymphocyte chromosomes can be separated into two fractions, each of which contains nucleoprotein. The fraction soluble in M NaCl consists largely of desoxyribose nucleohistone, and constitutes 90 to 92 per cent of the mass of the chromosome. The insoluble residue (the residual chromosome is a coiled thread containing some 12 to 14 per cent of ribose nucleic and about one-fifth as much desoxyribose nucleic acid; the residual chromosome accounts for 8 to 10 per cent of the mass of the chromosome. The staining of chromosomes—whether by the Feulgen procedure, by hematoxylin, orcein, or by basic dyes such as crystal violet—is due to the nucleohistone fraction which contains about 96 per cent of the nucleic acid of the chromosome. The form of the chromosome is due primarily to the protein thread of the residual chromosome. This thread is the only linear structure of microscopic dimensions in the chromosome that is not readily dispersed. When chromosomes are broken, it must be supposed that a break is made in the protein thread of the residual chromosome. The foregoing provides evidence for considering the residual chromosome to be the basis of the linear order of the genes. This would mean either that the residual chromosome is a structure around which the genes are organized or that the genes form part of its substance.     

THE CHROMOSOMES OF THE SEMPERVIVOIDEAE (CRASSULACEAE)

Uhl , Charles H. (Cornell U., Ithaca, New York.) Chromosomes of the Sempervivoideae (Crassulaceae). Amer. Jour. Bot. 48(2): 114–123. Illus. 1961.—Chromosome numbers are reported for 207 collections representing 68 of the ca. 95 species in this subfamily. Basic numbers are 16, 17, 18, and 19 in Sempervivum, Section Sempervivum (10 species, with many tetraploids and one hexaploid); 19 in Sempervivum, Section Jovisbarba (5 species, all diploid); 15 in Aichryson (9 species, including 1 aneuploid, 1 tetraploid, and 1 aneutetraploid); and strictly 18 in Aeonium (31 species, including 4 wholly and 1 partly tetraploid), Greenovia (3 species, 1 partly tetraploid), and Monanthes (10 species, including 2 wholly and 1 partly tetraploid). The cytological evidence appears decisive in ranking several species of disputed generic position definitely with Aichryson rather than with Aeonium. Possible relationships between various Canarian genera and certain North African species often classed in Sedum are discussed briefly in the light of the scanty morphological and cytological evidence. It is suggested that both these groups may be descended from the same ancestors that were widespread in North Africa before the deserts developed.     

THE STRANDEDNESS OF MEIOTIC CHROMOSOMES FROM ONCOPELTUS

Meiotic chromosomes were isolated from male Oncopeltus fasciatus by dissecting the testes under insect Ringer's solution and spreading the living cells on the Langmuir trough. After being dried by the critical point method, preparations were examined under the electron microscope. Chromosomes at all stages of prophase prove to be multistranded. A significant increase in the number of parallel 250 A fibers in the chromosomes occurs between zygotene and diakinesis. Parallel folding, rather than true multistrandedness, is interpreted as the mechanism responsible for this observed increase in multistrandedness. It has not been possible to determine whether the multistrandedness observed at leptotene represents true multistrandedness or is the result of parallel folding. Apparent multistrandedness is lost at metaphase when the 250 A fibers of the chromosomes become coiled more tightly. In preparations isolated by these methods, no structures other than the 250 A chromosome fibers are visible in the chromomeres, which appear as regionally coiled or folded areas of the fibers along the arm of the chromosome.     

THE CHROMOSOMES OF PARTHENOGENETIC FROGS AND TADPOLES

1. This paper presents cytological observations upon Dr. Loeb''s parthenogenetic frog material, with considerations upon the mechanism by which the diploid number and both sexes may be produced. 2. Both sexes of adults and tadpoles are present. 3. The chromosome number is diploid and probably 26 in both sexes. Sex chromosomes cannot be distinguished. 4. The chromosome numbers observed by other authors in parthenogenetic frog material are haploid, diploid, and variable. Their significance is considered. 5. The mechanism producing the diploid number, based on European observations, appears to be a doubling of the haploid number at some time after the second polar body is given off. 6. Overripeness may be a factor in producing both sexes of parthenogenetic frogs and tadpoles. 7. Genetic data indicate that the normal male is digametic and that there are differences of potency between male and female factors for sex which vary in frogs of the two races and in strains within the race. These differences have been interpreted by Witschi as forming a series of multiple allelomorphs.     

THE CHROMOSOMES OF PACHYPHYTUM (CRASSULACEAE)

Eleven of the 12 species of Pachyphytum, all that are available, have n = 31–33 standard chromosomes, or a multiple. Accessory chromosomes were found in some or all collections of four species; some cells of one plant have more than 50 of them. Accessory chromosomes often occur in groups at metaphase I, corresponding to their origin from one to several chromocenters of prophase I. Intraspecific polyploidy occurs within five species, with diploids to 12-ploids (n = ca. 186) in P. compactum and diploids to decaploids (n = ca. 160) in P. hookeri. Although the basic chromosome number is high, evidence from meiosis in certain hybrids shows that the basic 31–33 chromosomes are probably all different: they do not pair with each other and they do not duplicate each other. Polyploids, with 62 or more chromosomes, are probably autopolyploids: they form multivalents, and the chromosomes they contribute to hybrids pair with each other. Three different probable hybrids have been found in the wild, and more than 300 hybrids have been produced in cultivation.     

蜱类染色体的核型研究进展

总结了110种蜱的染色体核型。其中:软蜱科3属26种,包括锐缘蜱属12种,钝缘蜱属12种和残喙蝉属2种;硬蜱科8属84种,包括硬蜱属11种、牛蜱属3种、血蜱属16种、革蜱属12种、盲花蜱属5种、花蜱属19种、扇头蜱属6种和璃眼蜱属12种。 性别决定为XO,XX,XY,XX和X_1X_2Y,X_1X_1X_2X_23个系统。