the principles of heredity formulated by the Austrian Augustinian monk Gregor Mendel in 1865. These principles compose what is known as the system of particulate inheritance by units, or genes. The later discovery of chromosomes as the carriers of genetic units supported Mendel's two basic laws, known as the law of segregation and the law of independent assortment. In modern terms, the first of these states that genes are transferred as separate and distinct units from one generation to the next. The two members (alleles (allele)) of a gene pair, one on each of paired chromosomes, separate during the formation of sex cells by a parent organism. One-half of the sex cells will have one form of the gene, one-half the other form; the offspring that result from these sex cells will reflect those proportions. A modern formulation of the second law, the law of independent assortment, is that the alleles of a gene pair located on one pair of chromosomes are inherited independently of the alleles of a gene pair located on another chromosome pair and that the sex cells containing various assortments of these genes fuse at random with the sex cells produced by the other parent.