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4.1 hybrids: offspring that differ from their parents in one or more traits. Interspecific hybrids result from the union of two different species. Early Beliefs and Mendel The idea that biological traits are inherited existed long before the mechanisms of inheritance and gene interaction were understood. Stone tablets crafted by the Babylonians 6000 years ago show the pedigrees of successive generations of champion horses. Other carvings from the same period show the artificial crosspollination of date palms. Early records kept by Chinese farmers provide evidence of methods used for improving different varieties of rice. The selection of desired traits was based on keen observation and, to a great extent, trial and error. Early naturalists made assumptions about some incredible cross-species hybrids. The giraffe, for example, was thought to have been a cross between a leopard and a camel. (The fact that camels and leopards are not compatible did not seem to deter the theory.) The banana was thought to have been a hybrid of the acacia and the palm. Humans also mated selectively to produce desirable traits. The ancient Egyptians encouraged the intermarriage of royalty to preserve bloodlines. For example, Cleopatra married her younger brother. Plato, a Greek philosopher of the early 4th century B.C., called for the segregated mating of the elite. To maintain a line of strong warriors, the ancient Spartans practised infanticide, killing babies with undesirable characteristics. Pioneer of Genetics: Gregor Mendel Figure 1 Gregor Mendel was an Austrian monk whose experiments with garden peas laid the foundation for the science of genetics. One of the classic scientific experiments on inheritance was performed by an Austrian monk named Gregor Mendel (1822–1884) during the mid-19th century (Figure 1). Mendel’s work with garden peas not only explained the mechanism of gene inheritance in plants, but provided a basis for understanding heredity in general. When Mendel’s work was rediscovered many years later, it provided the missing piece in the theory of how organisms survive and reproduce. Why did Mendel choose the garden pea on which to perform his work? First, he observed that garden peas have a number of characteristics that are expressed in one of two ways (see Figure 2). For example, some garden peas produce green characteristics dominant trait recessive trait round wrinkled yellow green inflated constricted green yellow purple white seed shape dominant trait recessive trait side of stem end of stem tall short flower position seed colour pod shape pod colour Figure 2 The seven characteristics Mendel studied in his experiments with garden peas. Flower colour and seed colour are correlated. Plants with white flowers produce seeds that are yellow, and plants with violet-purple flowers produce seeds that are green. 130 Chapter 4 stem length flower colour 4.1 seeds (peas), while others produce yellow seeds. Some plants are tall, while others are short. Mendel also noticed different flower positions on the stem and different flower colours. The fact that there were only two ways for each trait to be expressed would make it easy to see which traits had been inherited from generation to generation. A second reason for using garden peas is the way the plant reproduces. Garden peas are both self-fertilizing and cross-fertilizing. Fertilization occurs when pollen produced by the stamen, the male part of the plant, attaches to the pistil, the female part (Figure 3). The pistil consists of the stigma, style, and ovary. The pollen grains fertilize the egg cells in the ovary. This process is called pollination. In self-fertilization, pollination occurs within one flower and the traits of the offspring are easily predicted. Mendel cross-pollinated the pea plants rather than allowing them to self-pollinate. He made sure to use purebreeding plants, that is, plants that always produce identical offspring. For example, tall plants produce only tall plants. If any offspring in any generation was not tall, then Mendel did not consider the parent plant to be pure and did not use it in his experiment. He transferred the pollen from one plant to the pistil of another plant, thus combining the male and female sex cells of different plants. To ensure that the recipient plant didn’t pollinate itself, he first removed its anthers. The pollen present then had to originate from the donor plant, resulting in seeds produced from cross-pollinated plants (Figure 4). stamen filament style anther ovary pollen stigma pistil Figure 3 The structure of a flower Mendel’s Experiments Mendel’s predecessors had hypothesized that the crossing of different traits would create a blend. According to this theory, crossing a plant that produced round seeds with one that produced wrinkled seeds would result in slightly wrinkled seeds. However, Mendel proved that this was not the case. When he crossed the pollen from a plant that produced round seeds with the eggs of one that produced wrinkled seeds, the offspring were always round. Did this mean that the pollen determines the seed coat? To test this idea, Mendel crossed the pollen from a wrinkled seed plant with the eggs from a round seed plant. Once again, all the offspring were round. In fact, the round trait dominated, regardless of whether the trait came from the male (pollen) parent or the female (seed) parent. Mendel repeated the procedure for other characteristics. He discovered that one trait always dominated another, whether the sex cell came from the male or female part of the plant. Tall plants produced tall offspring when cross-pollinated with short plants; likewise, plants that had yellow seeds produced offspring with yellow seeds when cross-pollinated with plants that had green seeds. Mendel reasoned that things called factors control the traits of a plant. The factors were later transfer pollen from pollen parent to seed parent remove anthers from seed parent Figure 4 The donor plant is also known as the pollen parent and the recipient is also known as the seed parent. Genes and Heredity 131 alleles: two or more alternate forms of a gene. The alleles are located at the same position on one of the pairs of homologous chromosomes. dominant: alleles of this type determine the expression of the genetic trait in offspring recessive: alleles of this type are overruled by dominant alleles, which determine the genetic trait called genes. He assumed that the genes control the inheritance of particular traits, such as seed colour and plant stem height. He also realized that there are alternate forms of a gene. Today, the alternate forms of a gene are called alleles. Green and yellow are expressions of the different alleles for seed colour. Tall stems and short stems are expressions of the different alleles for stem height. In garden peas, the traits that were expressed most often were considered to be dominant and those expressed less frequently were recessive. The allele for a yellow seed is dominant over the allele for a green seed; the allele for tall stems is dominant over the allele for short stems. Mendel cross-pollinated many plants and kept track of all the results. For each type of cross, he recorded the number of offspring that exhibited the dominant trait versus the recessive trait. He created a system of symbols to show what traits were passed to offspring. In this system, letters are used to represent traits. Uppercase letters stand for dominant traits, and lowercase letters stand for recessive traits. For the dominant trait of yellow seeds, Y represents the allele for yellow seeds; y represents the allele for green seeds, the recessive trait. Today, Mendel’s system is still in use. Mendel continued his experimentation by crossing two hybrid plants with round seeds from the first generation. He referred to the first generation as filial generation one, or F1 generation. The word filial comes from the Latin for son. Both of these F1 plants contain R and r alleles, one from each of their parents. This makes them hybrids. Remember, the R represents the round allele, while the r represents the wrinkled allele. You might predict an equal number of round and wrinkled offspring in the second, or F2, generation. However, this is not the ratio Mendel discovered when the two hybrids were crossed. He was astonished to find that 75% of the offspring expressed the dominant round trait, while only 25% expressed the wrinkled trait. How can these results be explained? Figure 5 shows what happens when the sex cells, or gametes, from the F1 generation recombine to form an F2 generation. All members of the F1 generation are round, but wrinkled offspring appear during the F2 generation. Any members of the F2 generation with an R allele will be round because the round allele is dominant over the wrinkled allele. To be wrinkled, the offspring must Rr Meiosis occurs. Each gamete has one of the homologous chromosomes. Figure 5 The result of crossing two hybrid pea plants with round seeds from the first generation 132 Chapter 4 R Rr r R r RR Rr Rr rr round round round wrinkled F2 generation inherits alleles from the gametes of the F1 generation. 4.1 Try This Activity Creating a Personal Profile Table 1 lists human traits controlled by dominant and recessive alleles. Table 1 Trait eye colour hair colour hairline freckles earlobe hair texture eyesight eyelashes nose line fingers Rh blood factor ear rim thumb joint finger hair folded hands tongue rolling clenched fist chin dimple blood type eyes Dominant brown or black or green brown or black pointed on forehead present suspended curly near or farsighted long convex tip 6 fingers positive Rh factor curled rim last joint bends out present left thumb over right can be rolled into U shape two wrist cords dimple in middle type A, B, AB astigmatism Recessive blue or grey blonde or red straight across forehead absent attached to head straight normal vision short concave or straight 5 fingers negative Rh factor not curled rim last joint is straight absent right thumb over left cannot be rolled three wrist cords no dimple type O no astigmatism Copy Table 2. Use information in Table 1 to complete your personal profile. Which additional traits can you include? Table 2 Trait (use the letter indicated) eye colour E/e hairline L/l earlobe T/t ear rim R/r freckles F/f thumb joint J/j finger hair P/p tongue rolling Y/y folded hands D/d nose line N/n hair colour H/h chin dimple G/g clenched fist K/k Appearance or Dominant or physical condition recessive ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? Possible genetic makeup ? ? ? ? ? ? ? ? ? ? ? ? ? Genes and Heredity 133