442 READINGS IN EVOLUTION, GENETICS, AND EUGENICS 



linkage with each other, A and C show a different and likewise fairly- 

 constant linkage strength, and so on through the entire group. This 

 leads to the conclusion that the genes of a linkage system are bound 

 together, gene with gene, with bonds of definite strength in each case. 

 In order to visualize the matter and get a more objective view of link- 

 age relations, Morgan and his associates have developed the chromo- 

 some theory of linkage. Its essential parts are: 



1. Genes which show linkage with each other are located in the 

 same pair of chromosomes. It is the substance of the chromosome 

 which binds the genes to each other and causes A to be inherited 

 when B is. 



2. Genes close together in the same chromosome show strong link- 

 age, genes farther apart show less linkage. 



3. Homologous chromosomes, those containing corresponding sets 

 of genes, one set derived from the father, one from the mother, lie side 

 by side (in synapsis) previous to the formation of gametes. At this 

 time breaks are likely to occur in the chromosomes and parts of one 

 are likely to replace corresponding parts of the other. 



4. Such replacement is called crossing-over. 



5. Breaks are commoner in long chromosomes than in short ones, 

 and between distant points than between near points on the same 

 chromosome. 



6. The genes occur in a chromosome, like beads on a string, in a 

 single row and in definite order. 



The supposed order of the genes in the four linkage groups of Dro- 

 sophila and their relative distances apart are shown in Fig. 90 (p. 437) . 

 In these diagrams or "maps," when the probable order of the genes 

 in a system has once been determined, the supposed end gene of the 

 system is placed at position o and the gene next to it is placed at a 

 distance (in centimeters or other units) corresponding to the average 

 cross-over percentage between the two, this process being repeated 

 from gene to gene until the whole chain is plotted. The "map" is 

 thus based on a summation of the distances (measured in cross-over per- 

 centages) from gene to gene. But if we compare the "map distances" 

 between genes not adjacent to each other in the chain with the observed 

 cross-over percentages between the same genes, we find that the map 

 distance is regularly greater than the cross-over percentage, except for 

 very short distances (5 or less). Thus if three genes occur in the order 

 ABC, it is usually found that AB+BC is greater than AC. In other 

 words, the cross-over percentage between B and C is commonly 



