Mendelian Inheritance

Transcription

1 Principles of Biology 38 contents Mendelian Inheritance Inheritance follows the rules of probability. Dice and the law of probability. Predicting the likelihood of inheriting a particular trait is similar to predicting the possible outcomes of rolling dice. It is important to understand probability to study genetic inheritance. Adrienne Hart-Davis/Science Source. Topics Covered in this Module Laws of Probability The Punnett Square Major Objectives of this Module Analyze a basic Punnett square problem. Analyze a test cross problem. Determine probabilities in monohybrid and dihybrid crosses. page 198 of pages left in this module

2 Principles of Biology contents 38 Mendelian Inheritance Laws of Probability Probability examines how likely it is that an event will occur. Probabilities range from 0, where an event has absolutely no chance of occurring, to 1, where there is no chance that the event will not occur. There are two basic rules to calculating probabilities depending on the type of events. Multiplication rule: Predicts the probability of independent events (a particular event in one experiment does not affect the probability of event in a second experiment). For example, when rolling a dice twice, the outcome in the first roll will not affect the probable outcomes when the dice is rolled a second time. The results of the two dice rolls are independent of each other. Each individual probability is multiplied to obtain an overall probability. Addition rule: Predicts the probability of mutually exclusive events. Mutually exclusive means that if one event occurs (such as a flipped coin comes up as a heads), the other event cannot simultaneously occur (i.e., it cannot be tails). The probability of either of the mutually exclusive events, either A or B, occurring is equal to the probability of event A added to the probability of event B (in the simple coin tossing case, the additions rule tells us that the probability that a flipped coin will be either heads or tails is 100%, because it will turn up heads approximately half of the time and tails the other half). Gregor Mendel studied the patterns of inheritance by breeding pea plants. In one set of experiments, he bred lines of plants that each had a different trait (for example, seed color or shape). He then crossed those lines with different traits to create a monohybrid cross; the offspring (F 1 ) did not have all of the traits that he observed in the parent generation. After crossing individuals in this F 1 generation, the traits that had disappeared in the F 1 generation reappeared in the offspring of these crosses (F 2 ). However, the trait did not reappear in all of the offspring on average, it appeared in 1 of 4 plants. He called this trait recessive because it could be masked by the other, dominant trait. What if Mendel had bred only four plants instead of hundreds? Would he have seen this pattern? What does probability have to do with genetics? The products of a genetic cross can be predicted using laws of probability. Mendel's principle of segregation states that two alleles in a parent segregate to form gametes with only one of those alleles through meiosis (Figure 1). The discovery of chromosomes in the latter part of the nineteenth century by Walther Flemming, along with the identification of the cell division process that gives rise to gametes (meiosis), reestablished the importance of Mendel's work. In meiosis, the parent reproductive cell, which contains a pair of each chromosome (called homologous chromosomes), undergoes two steps of division (meiosis I and meiosis II). Each parent cell divides into two cells during meiosis I, and the homologous chromosomes separate, followed by a second division in meiosis II where the two sister chromatids separate, forming four gametes that all contain a single allele (Figure 1). When alleles are described as dominant or recessive, it is important to keep in mind that dominance and recessiveness are not intrinsic allelic properties. Instead, they are effects that may only be measured in relation to the effects of the alternative alleles of the same gene.

3 Figure 1: Mendel's principle of segregation. Homologous chromosomes contain alternative versions of the same allele. In this case, the blue chromosomes contain the allele "G" or "g," and the red chromosomes contain either "W" or "w." Note how the homologous chromosomes and their sister chromatids randomly segregate into the four different gametes based on their positions during meiosis I, leading to the two alternative pathways. The properties of the allele (dominant or recessive) do not influence the location of the alleles in gametes Nature Education All rights reserved. Figure Detail Which allele ends up in which gamete is random. However, one could predict the likelihood of the offspring possessing specific alleles (the genotype), and ultimately the phenotype, using the rules of inheritance that are governed by probability. Probability examines how likely it is that an event will occur. BIOSKILL What Is the Probability of a Coin Toss? Every time one flips a standard (fair) coin, the chance of getting heads is 1 in 2, or 50% (Figure 2). Each toss is one independent probability event the outcome of one toss does not impact the outcome of any other tosses. You likely understand the reasoning behind the concept of independent probability, but it is not so easy to observe. If one flips a coin twice, there is a good chance that it may be heads both times. If one flips a coin 10 times, there is a chance, but a much smaller chance, that it will be heads all 10 times. During World War II, a South African statistician, John Kerrich, passed some of the time he spent as a prisoner of war flipping a coin 10,000 times.

4 He ended up with heads 50.67% of the time. The more events there are, the more the results reflect the predicted probabilities. Figure 2: Calculating the probability of several independent events. The total probability of multiple independent events is calculated by first determining the probability of one of the independent events. In this case, the probability of a single coin toss resulting in heads is one-half, or 50%. Therefore, the total probability of four coin tosses resulting in heads is ½ ½ ½ ½, or one-half to the fourth power Nature Education All rights reserved. Figure Detail Consider a person flipping a coin four times. What is the chance of getting heads on all four tries? Each of the four tries has a probability of 1/2 that it will be heads. Since none of the flips will impact the outcome of any of the other flips, you multiply the probability of heads for each flip together to determine the probability of four heads in a row. ½ x ½ x ½ x ½ = (½) 4 = 1/16 = = 6.25% There is a bit more than a 6% chance of getting heads on all four coin flips. This example demonstrates the multiplication rule for independent probability. BIOSKILL How does the multiplication rule apply to genetics? A recessive trait is expressed only when the allele coding for the recessive trait is inherited from both parents (that is, the organism is homozygous for the recessive trait). If the recessive allele is only inherited from one parent (i.e., the organism is heterozygous), the dominant trait is expressed. In Mendel's pea plants, white flowers were recessive, and purple flowers were dominant. Mendel reasoned that if each parent carried the alleles coding for one dominant and one recessive trait, the offspring had a 50% chance of inheriting the allele coding for the recessive trait from one parent and a 50% chance of inheriting the allele coding for the recessive trait from the other parent.

5 Test Yourself Imagine that two heterozygous plants, each with one dominant allele and one recessive allele, are mated with each other. Since a plant has a 50% chance of inheriting a particular allele from each parent, what is the probability that the offspring of the two plants will inherit two copies of the allele coding for the recessive trait? Submit There is a 25% chance that such a plant will have white flowers and, conversely, a 75% chance that it will have purple flowers. The reason that white flowers, the recessive trait, are less likely is that the recessive trait can only occur under one scenario: namely, the offspring's inheritance of both recessive alleles; conversely, there is a greater chance that the offspring will have purple flowers because purple flowers can result in two different ways: plants that are homozygous for the dominant trait and those that are heterozygous both express the purple flower trait. As a result, when counting plants with different flower colors, we expect that purple flowers should occur in a 3:1 ratio with white flowers. Mendel wanted to test this hypothetical ratio, but he knew that patterns in probability are most reliable when the numbers are large. As a result, he measured hundreds of offspring from multiple monohybrid crosses over many generations of pea plants. IN THIS MODULE Laws of Probability The Punnett Square Summary Test Your Knowledge PRIMARY LITERATURE Using sterile mates and engineered toxins to beat bugs Suppressing resistance to Bt cotton with sterile insect releases. View Download Classic paper: How scientists cloned the first mammal (1997) Viable offspring derived from fetal and adult mammalian cells. View Download Classic paper: Fruit fly research reveals how complex organisms form (1980) Mutations affecting segment number and polarity in Drosophila. View Download SCIENCE ON THE WEB Genetics in the Lab Simulate a laboratory to discover the genetic inheritance patterns of fruit flies Let the Computer Do the Work Use this website to compare observed and expected frequencies using the chi-square test Roll the Dice Observe the odds of rolling a certain number Model Organism Explore this NIH website about model organisms used in biomedical research In Mendel's Words Read Gregor Mendel's original paper Experiments in Plant Hybridization (1865) Understanding the Spectrum of Dominance

6 Alleles are never exclusively "dominant" or "recessive." page 199 of pages left in this module

7 Principles of Biology contents 38 Mendelian Inheritance The Punnett Square The Punnett square, invented by geneticist Reginald Punnett, is used to assess probability in genetics (Figure 3). The father's alleles are listed on one side of the square and the mother's on the other side. The cells of the table represent offspring, each of which receives one allele from the father and one from the mother. Drawing a Punnett square is one way to figure out the probability of parents with particular genotypes having offspring of a certain genotype or phenotype. Punnett squares are often easier to visualize than mathematical calculations of probability. Illustrating probability and the Punnett square with Mendel's peas. Not all pea plants with purple flowers have the same set of alleles. A purpleflowered plant may be homozygous, having inherited the dominant allele from each parent, or heterozygous, having inherited one dominant and one recessive allele. What is the probability that a plant with heterozygous parents will also be heterozygous? A plant is heterozygous when it receives or an allele coding for the dominant trait from the female and an allele coding for the recessive trait from the male (event A) an allele coding for the dominant trait from the male and an allele coding for the recessive trait from the female (event B). These two possibilities are not independent in fact, they are mutually exclusive. In such cases, probabilities are calculated using the addition rule. What is the probability that one of these events will occur? In this case, the two events have the same likelihood of occurring. A monohybrid cross between two heterozygotes gives four offspring. The probability of event A is one-quarter, and the probability of event B is one-quarter. Applying the addition rule to this situation, we would add one-quarter to one-quarter to conclude that there is a 50% chance (¼ + ¼ = ½ ) that one of the offspring will be heterozygous. Thus, if two heterozygotes breed, half of their offspring, on average, will also be heterozygotes. Because purple flowers are dominant, this phenotype does not reveal the plant's genotype. What is the probability that a plant is heterozygous if its parents are heterozygotes with purple flowers? The probability that a plant is heterozygous, given that it has purple flowers, is two-thirds. Note that this is higher than the chance of the plant being heterozygous when its phenotype is unknown, which we calculated to be one-half. An easier way to visualize these probabilities is by taking the alleles of each parent and combining them by means of a Punnett square (Figure 3).

8 Figure 3: Probability predicts genotype and phenotype in Mendel's monohybrid cross. Probability is used to assess the genetic outcomes (both genotype and phenotype of the offspring). This Punnett square is a straightforward way to visualize the possible combinations of alleles the offspring of two parents will have. In this example, two individuals that are heterozygous (Pp) are mated. The alleles in the parental gametes are on the outside of the square, and each cell of the Punnett square represents the possible allelic combinations for the offspring. (Each offspring, of course, receives one allele from the male's gametes (sperm) and one allele from the female's gametes (egg).) The cells show that on average three of every four offspring should have purple flowers; on average, two out of those three will be heterozygous (Pp), and one out of three will be homozygous for the dominant trait (PP). Only one of four offspring, on average, will have white flowers as a result of being homozygous recessive (pp) Nature Education All rights reserved. How do I use a Punnett square? The body of a fruit fly may be black or brown. Brown is the dominant color (B) and black is the recessive color (b). You can create a Punnett square to figure out what the offspring will look like given the parents' genotypes

9 (Figure 4). Figure 4: How to draw a Punnett square. Punnett squares may predict the traits of the offspring that result from a particular cross Nature Education All rights reserved. Figure Detail The below instructions describe how to create a Punnett square for a cross between a homozygous brown female fly (BB) and a homozygous black male fly (bb). These instructions could also be used for monohybrid and dihybrid crosses. Larger crosses, however, are more complicated. Step 1. Write down the genotype of the female and male. In this case, the female's genotype is BB, and the male's is bb. Count each parent's alleles and multiply the numbers. The result determines how big the Punnett square needs to be. In this case, because there are only two alleles for each parent, there are four possible outcomes of allelic combinations (2 * 2 = 4), and a square with 4 cells is needed. Step 2. Draw a square with the required number of cells. Step 3. List the female's alleles along one side of the square and the male's alleles along the other. In this case, the alleles from the female are written in the column on the left (B and B) and the alleles from the male are written in the row on the top (b and b). Step 4. Copy each combination of alleles that can be received from each

10 parent in the respective cells. Each B from the female is combined with each b from the male in each cell, thereby providing the possible genotypes of the offspring. Step 5. Determine the phenotypes of the offspring based on their genotypes. In this case, all of the individuals are heterozygous (Bb), and because the allele for brown (B) is dominant over the allele for black (b), all the fruit flies of this pairing would be brown. BIOSKILL Crossing Flies: Practicing the Punnett Square Practice determining phenotypes and phenotypic ratios expected from the offspring (F 1 generation) and the offspring's offspring (F 2 generation) of a monohybrid cross between a brown female with dominant homozygous alleles (BB) and a black male with recessive homozygous alleles (bb) (Figure 5). Figure 5: Monohybrid crosses. Determine the phenotypes and phenotypic ratio expected from the offspring (F 1 generation) and the offspring's offspring (F 2 generation) of a monohybrid cross between a brown female with dominant homozygous alleles (BB) and a black male with recessive homozygous alleles (bb) Nature Education All rights reserved. Transcript Test Yourself What are the possible genotypes that could produce a brown phenotype? What about a black phenotype? Submit

11 BIOSKILL Using Punnett squares for dihybrid crosses. The F 2 generation in the interactive in Figure 5 represented a monohybrid cross, which is a pairing of two parents that are both heterozygous for one trait (in this case body color). What if the parents differ not just in one trait but in two or more? Fruit flies may have brown or black bodies and brown or red eyes. A dihybrid cross is a cross between parents that are heterozygous for two traits. Although a Punnett square for a dihybrid cross is more complex than Punnett squares for monohybrid crosses, the purpose of the square is the same: determining the probable genotypes and phenotypes of the offspring. Figure 6: Dihybrid Cross. When both parents are heterozygous for two traits, the offspring may have any of four possible phenotypes (brown body and red eyes, brown body and brown eyes, black body and red eyes, and black body and brown eyes). Note that allele combinations for a dihybrid cross are made by determining the possible combinations of alleles from the two different genes Nature Education All rights reserved. Figure Detail

12 Test Yourself What is the phenotypic ratio among the offspring of the dihybrid cross shown in Figure 6? Submit What is a test cross? In Mendel's peas, the purple trait is dominant, so a purple flower could either be homozygous (PP) or heterozygous (Pp) for the purple allele. How could you experimentally determine the genotype if you had a pea plant and all that you knew about it was that it had a purple flower? In a test cross, the purple flower, whose genotype is unknown, is bred with a white flowered pea plant that expresses the recessive trait (Figures 7 and 8). If the purple plant is homozygous, all of the offspring will be purple because they will all have at least one copy of the dominant purple allele. If the parent plant is heterozygous, having both an allele coding for the dominant trait and an allele coding for the recessive trait, half of the offspring will be white and the other half will be purple. Therefore, you could determine whether the plant with an unknown genotype is heterozygous or homozygous by looking at the phenotypes of the offspring from a test cross. Figure 7: Test cross. Crossing an organism of unknown genotype with one expressing recessive traits will determine the genotype of the unknown parent (i.e., whether it is homozygous or heterozygous) Nature Education All rights reserved. Figure Detail

13 Figure 8: Test-cross problem. What is a brown female parent's genotype if half of her offspring are brown (B), the other half are black (b), the black allele is recessive, and the male parent is black (bb)? 2013 Nature Education All rights reserved. Test Yourself In a cross between a brown female fly and a black male fruit fly, if half of the offspring are black, what is the female's genotype? What if all of the offspring are brown? Submit BIOSKILL A Chi-Square Test Uses Probability to Detect Differences between Expected and Observed Values The predictions made in Mendel's monohybrid and dihybrid cross are based on some assumptions that certain events are random and will occur with a certain probability. For example, it assumes that alleles randomly and independently assort into gametes and that fertilization between gametes is random. Therefore, the simple predictions from Mendelian genetics are not always observed. If the observed data differ substantially from what would have been expected from our Punnett square and probability calculations, we can infer that the simple assumptions, such as independent assortment, are not true. When we assume that the observed data of a genetic cross will fit a given Mendelian ratio, a null hypothesis (H 0 ) is generated, which is that there will be no real difference between the measured values (or ratio) and the expected values (or ratio). The null hypothesis can either be rejected or not rejected for a given set of data measured using statistical analysis. One of the simplest statistical tests for assessing whether the observed values differ substantially from those expected is the chi-square test, often abbreviated χ 2. In figure 9, a chi-square test is illustrated using a cross between genes for body color and wing type in fruit flies. In these genes, brown bodies (B) and normal wings (W) are dominant traits, and black bodies (b) and vestigial wings (w) are recessive traits. A female fruit fly heterozygous for brown body and normal wings (BbWw) was crossed with a male fruit fly with a black body and vestigial wings (bbww) (Figure 9).

14 Figure 9: Crossing a heterozygous dominant female with a homozygous recessive male. Four equally probable phenotypes are expected in the offspring Nature Education All rights reserved. If the assumptions are that these two genes are independent and randomly assort into gametes, we can predict the proportions of genotypes and phenotypes using a Punnett square as above. In this cross, we expect a 1:1:1:1 ratio of the following phenotypes: brown body, normal wings; black body, vestigial wings; black body, normal wings; and brown body, vestigial wings. That is, on average, one-quarter (25%) of the offspring from this cross should be in each of the four phenotype categories. If a scientist breeds 2,300 fruit flies, each phenotype should be represented by approximately 575 flies (one-quarter expected in each group x 2,300 flies = 575 flies per group). Remember that it may take many crosses before a reliable estimate of proportions can be calculated (if the scientist only bred five or 10 individuals, the proportions would almost certainly differ from the expected ratios, but this could have easily happened by chance). However, in this case, the observed results look rather different from the results that would be expected from random processes (Figure 10). Are these results significantly different than expected? Figure 10: Data for χ 2 analysis. A chi-square analysis is used to determine if the observed counts are significantly different from the expected counts for each of these four fruit fly phenotypes.

15 2011 Nature Education All rights reserved. This is where the chi-square test comes in. The formula is χ 2 = the sum of ([observed count - expected count] 2 /expected count) If the observed counts were exactly as expected, the χ 2 value will be 0. As the observed results increasingly differ from the expected results, the χ 2 value increases. To interpret the χ 2 result, the degrees of freedom (df) must be also determined, which are equal to "n-1," where "n" is the number of different categories into which the data are divided. In the example above, there are 4 possible phenotypes, and thus "n" is 4; the df is 3 (4-1). A χ 2 probability table is used to determine the statistical significance of the χ 2 value we calculate; that is, the table tells us what the probability is that our observed values differ from the expected value by chance alone. A high probability (p), such as p > 0.5, tells us that there is more than a 50% chance that the differences that we found between the observed and the expected results could have been simply due to chance. This would not give us much confidence that the results were in fact significantly different and we would not reject the null hypothesis that any observed differences were simply due to chance. To find the significance value, match the df and χ 2 values in a χ 2 probability distribution table (available in most statistical textbooks or online). One common statistical standard is p < 0.05, which means that there is less than a 5% probability that the observed results would have occurred simply by chance and were due to random sampling; as a corollary, there is a greater than 95% probability that the observed results are due to non-random factors, and that the null hypothesis can be rejected. For these fruit fly crosses, χ 2 = 1,002.65, and df = 3. For three degrees of freedom, the χ 2 value that must be found for there to be only a 5% that the differences are due to chance (p < 0.05) is That is, any χ 2 value greater than 7.82 has less than a 5% probability of occurring by chance. Our χ 2 value of more than 1,000 is much greater than 7.82! In other words, this result is highly unlikely to have occurred by chance. What could account for it? Observed phenotypic ratios often differ from those expected by Mendelian inheritance. This is because many genes are not inherited according to Mendel's three principles of inheritance (dominance, segregation and independent assortment). For example, some genes do not code for clear dominant or recessive traits, violating the principle of dominance. In addition, genes that are near each other on the same chromosome are sometimes linked so that they are not sorted independently into gametes, violating the principle of independent assortment. Although Mendelian inheritance only applies to a minority of genes, Mendel's work pioneered the quantitative genetic methods that allowed the discovery of non-mendelian traits. BIOSKILL BIOSKILL What Makes a Suitable Model Organism? A model organism is one that is well suited to scientific research. Experiments using model organisms help scientists understand the fundamentals of biological principles, which can then be extended to other organisms, including humans. Gregor Mendel used pea plants as a model organism in the mid-1800s, and in the early 1900s Thomas Morgan used the fruit fly Drosophila melanogaster. Other organisms now commonly used in genetic research include Caenorhabditis elegans (a nematode worm), Arabidopsis thaliana (a small plant in the mustard family), mice and yeast. What makes these organisms useful as models? Pea plants are inexpensive and easy to grow from seeds. They mature quickly, produce many seeds, and show a number of visible

16 variations in flower color, seed color and shape. Gregor Mendel also cut the flowers' male reproductive appendages, or stamens, to only allow cross-pollination. This allowed the parents of the offspring to be known and traced. Today, the most common plant used as a model organism is Arabidopsis thaliana, which shares some of the same advantages as pea plants, such as high variability in traits, but is also much smaller and can grow more rapidly. It also has a small genome with relatively few chromosomes, making modern genetic analyses more streamlined. Drosophila melanogaster and Caenorhabditis elegans offer many of the same benefits described above. These invertebrates are small and easy to raise in a lab. Both lay many eggs, and their eggs mature quickly, allowing many generations of inheritance to be observed over a short period of time. Drosophila, in particular, have many visible variable characteristics, such as wing type, body color and eye color (as mentioned above), which allow their genotypes to be traced. Fruit flies also have relatively small genomes with only four pairs of chromosomes. Mice are mammals and share many characteristics with humans. In fact, about 75% of rodent genes share largely the same function as those in humans, making mice genetically closer to humans than any of the other common model organisms. Mice may be used to examine questions that researchers are interested in but that would be unethical to test using human subjects. The mouse genome is also useful in research because genes may be "knocked out," an experimental technique in which scientists inactivate a target gene or replace it with an inactivated mutant version to see how its absence affects the organism. The other model organisms may also have their genes knocked out to address important questions in gene function. The simplicity of baker's yeast (Saccharomyces cerevisiae), a eukaryotic organism that is unicellular, has made it useful for studying cell division and gene regulation how genes are turned "on" and "off." It is the easiest eukaryotic genome to manipulate because it is so simple compared to other eukaryotes, but its genes still share many of the basic functions of more complex eukaryotes. This helps scientists identify how genes work on a cellular level in yeast, and apply that understanding to organisms like humans. Test Yourself Why have so many genetic experiments been done on fruit flies? Name two characters that make fruit flies a good model organism. Submit BIOSKILL IN THIS MODULE Laws of Probability The Punnett Square Summary Test Your Knowledge page 200 of pages left in this module

17 Principles of Biology contents 38 Mendelian Inheritance Summary OBJECTIVE Analyze a basic Punnett square problem. Punnett squares are a way to visualize the probabilities of genotypes and phenotypes in offspring using parental genotypes. For example, using a Punnett square to analyze a monohybrid cross, a phenotypic ratio of 3:1 dominant to recessive in offspring may be predicted. In this case, one-third of the offspring with the dominant phenotype are homozygous, and two-thirds are heterozygous for the trait. OBJECTIVE Analyze a test cross problem. To identify the unknown genotype of an organism showing the dominant phenotype, a test cross of an organism expressing the dominant phenotype is made with an organism displaying recessive traits. The distribution of phenotypes in the offspring will reveal whether the unknown parent genotype is heterozygous or homozygous. A chi-square test is used to evaluate whether the difference between the observed results and the expected results of a test cross is likely to have occurred by chance or whether it represents an inheritance pattern different from the one predicted. OBJECTIVE Determine probabilities in monohybrid and dihybrid crosses. The principle of independent assortment states that when gametes form, each pair of alleles assorts independently. Which offspring inherit which alleles is random, but we are able to predict the likelihood of phenotypes in the offspring based on the parents' genotypes and the rules of probability. In a monohybrid cross, the ratio of the phenotypes in the offspring is 3:1 dominant:recessive, or three-quarters dominant and one-quarter recessive. In a dihybrid cross, the ratio of phenotypes in the offspring is 9:3:3:1 (dominant, dominant):(dominant, recessive):(recessive, dominant): (recessive, recessive); stated differently, 9/16 of the offspring have both dominant phenotypes, 3/16 of the offspring have the dominant phenotype for gene 1 and the recessive phenotype for gene 2, 3/16 have the recessive phenotype for gene 1 and the dominant phenotype for gene 2, and 1/16 of the offspring have both recessive phenotypes. Key Terms addition rule Considering mutually exclusive events, the probability of both occurring is the sum of the probabilities of each event. heterozygous Having two different alleles for a trait. homozygous Having identical alleles for a trait. independent probability Probability of one event occurring has no effect on the probability of the other. model organism An organism that is well suited for scientific research for a variety of reasons. multiplication rule If the events are truly independent, the probability of both events happening is found by multiplying the probabilities of each event. null hypothesis A hypothesis assuming that there is no difference between the observed values

18 and the expected values. probability Calculation of the chance of an event occurring. Punnett square Table method used to assess probabilities of genotypic and phenotypic outcomes among genetic crosses. test cross A cross between an organism with a dominant phenotype but an unknown genotype and an individual that is homozygous recessive. IN THIS MODULE Laws of Probability The Punnett Square Summary Test Your Knowledge PRIMARY LITERATURE Using sterile mates and engineered toxins to beat bugs Suppressing resistance to Bt cotton with sterile insect releases. View Download Classic paper: How scientists cloned the first mammal (1997) Viable offspring derived from fetal and adult mammalian cells. View Download Classic paper: Fruit fly research reveals how complex organisms form (1980) Mutations affecting segment number and polarity in Drosophila. View Download SCIENCE ON THE WEB Genetics in the Lab Simulate a laboratory to discover the genetic inheritance patterns of fruit flies Let the Computer Do the Work Use this website to compare observed and expected frequencies using the chi-square test Roll the Dice Observe the odds of rolling a certain number Model Organism Explore this NIH website about model organisms used in biomedical research In Mendel's Words Read Gregor Mendel's original paper Experiments in Plant Hybridization (1865) Understanding the Spectrum of Dominance Alleles are never exclusively "dominant" or "recessive." page 201 of pages left in this module

19 Principles of Biology contents 38 Mendelian Inheritance Test Your Knowledge 1. Huntington's disease, a progressive nervous system disorder, is inherited as a dominant trait. The songwriter Woody Guthrie died of Huntington's disease. If he was heterozygous for the trait and his wife did not carry it, what was the probability that their son would develop Huntington's disease? 40% 25% 66% 10% 50% 2. If both parents are heterozygous for a trait, what is the probability that a given offspring of theirs will display the recessive trait? Assume the gene for the trait is not sex-linked. 2/3 3/4 1/2 1/4 1/3 3. In fruit flies, brown body color (B) is dominant to black body color (b). A brown fruit fly is crossed with a black fruit fly. Both flies are purebred. What will the offspring's genotypes be? 1/2 BB, 1/2 bb 1/4 bb, 3/4 Bb all Bb 1/4 bb, 1/2 Bb, 1/4 BB 1/3 BB, 2/3 Bb 4. In humans, dry earwax is recessive to wet earwax. Consider a woman with dry earwax and her male partner who has wet earwax. If he is heterozygous for the trait of wet earwax, what is the probability that their child will have wet earwax? This cannot be predicted based on the information provided. 50% 75% 25% 100% 5. In peas, yellow color and round shape are dominant to green color and wrinkled shape, which are recessive. If you have a round, yellow pea and want to know its genotype, what kind of pea should you breed it with? yellow-round yellow-wrinkled green-wrinkled green-round any type

20 Submit IN THIS MODULE Laws of Probability The Punnett Square Summary Test Your Knowledge PRIMARY LITERATURE Using sterile mates and engineered toxins to beat bugs Suppressing resistance to Bt cotton with sterile insect releases. View Download Classic paper: How scientists cloned the first mammal (1997) Viable offspring derived from fetal and adult mammalian cells. View Download Classic paper: Fruit fly research reveals how complex organisms form (1980) Mutations affecting segment number and polarity in Drosophila. View Download SCIENCE ON THE WEB Genetics in the Lab Simulate a laboratory to discover the genetic inheritance patterns of fruit flies Let the Computer Do the Work Use this website to compare observed and expected frequencies using the chi-square test Roll the Dice Observe the odds of rolling a certain number Model Organism Explore this NIH website about model organisms used in biomedical research In Mendel's Words Read Gregor Mendel's original paper Experiments in Plant Hybridization (1865) Understanding the Spectrum of Dominance Alleles are never exclusively "dominant" or "recessive." page 202 of 989