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The unique appearance of this white- skinned, blue-eyed alligator is the result of a genetic condition. SECTION 1 Mendel’s Legacy SECTION 2 Genetic Crosses Unit 5—Heredity Topics 1–6 CHAPTER 9 172 9 CHAPTER F UNDAMENTALS OF G ENETICS F UNDAMENTALS OF G ENETICS Copyright © by Holt, Rinehart and Winston. All rights reserved.
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CHAPTER 9 FUNDAMENTALS OF GENETICS - … OF GENETICS 177 MENDEL’S RESULTS AND CONCLUSIONS In one of his experiments, Mendel crossed a plant true-breeding for green pods with one

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Page 1: CHAPTER 9 FUNDAMENTALS OF GENETICS - … OF GENETICS 177 MENDEL’S RESULTS AND CONCLUSIONS In one of his experiments, Mendel crossed a plant true-breeding for green pods with one

The unique appearance of this white-skinned, blue-eyed alligator is the resultof a genetic condition.

SECTION 1 Mendel’s Legacy

SECTION 2 Genetic Crosses

Unit 5—HeredityTopics 1–6

C H A P T E R 9172

9CHAPTER FUNDAMENTALS OF

GENETICSFUNDAMENTALS OFGENETICS

Copyright © by Holt, Rinehart and Winston. All rights reserved.

Page 2: CHAPTER 9 FUNDAMENTALS OF GENETICS - … OF GENETICS 177 MENDEL’S RESULTS AND CONCLUSIONS In one of his experiments, Mendel crossed a plant true-breeding for green pods with one

173F U N D A M E N T A L S O F G E N E T I C S

M E N D E L’ S L E G A C YGenetics is the field of biology devoted to understanding

how characteristics are transmitted from parents to offspring.

Genetics was founded with the work of Gregor Johann Mendel.

This section describes Mendel’s experiments and the principles

of genetics that resulted from them.

GREGOR MENDELIn 1843, at the age of 21, Gregor Mendel, shown in Figure 9-1,entered a monastery in Brunn, Austria. His task of tending the gar-den gave him time to think and to observe the growth of manyplants. In 1851, he entered the University of Vienna to study sci-ence and mathematics. His mathematics courses included trainingin the then-new field of statistics. Mendel’s knowledge of statisticslater proved valuable in his research on heredity—the transmis-sion of characteristics from parents to offspring. When Mendelreturned to the monastery, he taught in a high school and alsokept a garden plot. Although he studied many plants, he is remem-bered most for his experiments with Pisum sativum, a species ofgarden peas.

Mendel’s Garden PeasMendel observed seven characteristics of pea plants. A characteris-tic is a heritable feature, such as flower color. Each characteristicoccurred in two contrasting traits. A trait is a genetically determinedvariant of a characteristic, such as yellow flower color. The peacharacteristics that Mendelobserved were plant height(traits: long and short), flowerposition along stem (traits:axial and terminal), pod color(traits: green and yellow), podappearance (traits: inflatedand constricted), seed texture(traits: round and wrinkled),seed color (traits: yellow andgreen), and flower color(traits: purple and white).Mendel used his knowledge of statistics to analyze hisobservations of these sevencharacteristics.

SECTION 1

O B J E C T I V E S● Describe how Mendel was able to

control how his pea plants werepollinated.

● Describe the steps in Mendel’sexperiments on true-breedinggarden peas.

● Distinguish between dominantand recessive traits.

● State two laws of heredity thatwere developed from Mendel’swork.

● Describe how Mendel’s results can be explained by scientificknowledge of genes andchromosomes.

V O C A B U L A R Ygeneticsheredity trait pollinationself-pollinationcross-pollination true-breedingP generation F1 generation F2 generation dominantrecessive law of segregation law of independent assortment molecular genetics allele

Copyright © by Holt, Rinehart and Winston. All rights reserved.

Gregor Johann Mendel lived from 1822to 1884. Mendel’s experiments withgarden peas led to his discovery of thebasic principles of genetics.

FIGURE 9-1

Page 3: CHAPTER 9 FUNDAMENTALS OF GENETICS - … OF GENETICS 177 MENDEL’S RESULTS AND CONCLUSIONS In one of his experiments, Mendel crossed a plant true-breeding for green pods with one

C H A P T E R 9174

Mendel collected seeds from his pea plants and carefullyrecorded each plant’s traits and seeds. The next year, he plantedthe seeds. He observed that purple-flowering plants grew frommost of the seeds obtained from purple-flowering plants but thatwhite-flowering plants grew from some of the seeds of purple-flowering plants. And when experimenting with the characteristicof plant height, he observed that while tall plants grew from mostof the seeds obtained from tall plants, short plants grew from someof the seeds obtained from tall plants. Mendel wanted to find anexplanation for such variations.

Mendel’s MethodsMendel was able to observe how traits were passed from one gen-eration to the next by carefully controlling how pea plants werepollinated. Pollination occurs when pollen grains produced in themale reproductive parts of a flower, called the anthers, are trans-ferred to the female reproductive part of a flower, called the stigma.

Self-pollination occurs when pollen is transferred from theanthers of a flower to the stigma of either that flower or anotherflower on the same plant. Cross-pollination occurs between flowersof two plants. Pea plants normally reproduce through self-pollination.

Self-pollination can be prevented by removing all of the anthersfrom the flowers of a plant. Then, cross pollination can be performedby manually transferring pollen from the flower of a second plant tothe stigma of the antherless plant, as Figure 9-2 shows. By prevent-ing self pollination and manually cross-pollinating pea plants,Mendel selected parent plants that had specific traits and observedthe traits that appeared in the offspring.

Copyright © by Holt, Rinehart and Winston. All rights reserved.

Mendel controlled the breeding of hispea plants and tracked the inheritanceof traits by transferring pollen from theanthers of one plant to the stigma ofanother plant.

FIGURE 9-2

www.scilinks.orgTopic: Gregor MendelKeyword: HM60698

Page 4: CHAPTER 9 FUNDAMENTALS OF GENETICS - … OF GENETICS 177 MENDEL’S RESULTS AND CONCLUSIONS In one of his experiments, Mendel crossed a plant true-breeding for green pods with one

1 2 3

175F U N D A M E N T A L S O F G E N E T I C S

MENDEL’S EXPERIMENTSMendel initially studied each characteristic and its contrasting traitsindividually. He began by growing plants that were true-breeding foreach trait. Plants that are true-breeding, or pure, for a trait alwaysproduce offspring with that trait when they self-pollinate. For exam-ple, pea plants that are true-breeding for the trait of yellow pods self-pollinate to produce offspring that have yellow pods. Mendelproduced true-breeding plants by self-pollinating the pea plants forseveral generations, as Figure 9-3 shows. He eventually obtained 14true-breeding plant types, one for each of the 14 traits observed.

Mendel cross-pollinated pairs of plants that were true-breedingfor contrasting traits of a single characteristic. He called the true-breeding parents the P generation. He cross-pollinated by trans-ferring pollen from the anthers of one plant to the stigma ofanother plant. For example, if he wanted to cross a plant that wastrue-breeding for the trait of yellow pods with one that was true-breeding for the trait of green pods, he first removed the anthersfrom the plant that produced green pods. Then, he dusted thepollen from a yellow-podded plant onto the stigma of a green-podded plant and allowed the seeds to develop.

When the plants matured, Mendel recorded the number of eachtype of offspring produced by each cross. He called the offspring ofthe P generation the first filial generation, or F1 generation. Hethen allowed the flowers from the F1 generation to self-pollinateand collected the seeds. Mendel called the plants in this generationthe second filial generation, or F2 generation. Following thisprocess, Mendel performed hundreds of crosses and documentedthe results of each by counting and recording the observed traitsof every cross. Table 9-1 summarizes the results of many ofMendel’s crosses.

Copyright © by Holt, Rinehart and Winston. All rights reserved.

filial

from the Latin filialis, meaning “of a son or daughter”

Word Roots and Origins

Mendel bred plants for severalgenerations that were true breeding forspecific traits. These plants he called theP generation. He then observed thepassage of these specific traits throughsuccessive generations.

FIGURE 9-3

Page 5: CHAPTER 9 FUNDAMENTALS OF GENETICS - … OF GENETICS 177 MENDEL’S RESULTS AND CONCLUSIONS In one of his experiments, Mendel crossed a plant true-breeding for green pods with one

Copyright © by Holt, Rinehart and Winston. All rights reserved.

P generation

axial ! terminal

tall ! short

inflated ! constricted

green ! yellow

round ! wrinkled

yellow ! green

purple ! white

F1 generation

all axial

all tall

all inflated

all green

all round

all yellow

all purple

F2 generation

651 axial207 terminal

787 tall277 short

882 inflated299 constricted

428 green152 yellow

5,474 round1,850 wrinkled

6,022 yellow2,001 green

705 purple224 white

Observedratio

3.14:1

2.84:1

2.95:1

2.82:1

2.96:1

3.01:1

3.15:1

Predictedratio

3:1

3:1

3:1

3:1

3:1

3:1

3:1

C H A P T E R 9176

TABLE 9-1 Mendel’s Crosses and Results

Characteristic

Position of flowers along stem

Height of plant

Pod appearance

Pod color

Seed texture

Seed color

Flower color

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177F U N D A M E N T A L S O F G E N E T I C S

MENDEL’S RESULTS AND CONCLUSIONS

In one of his experiments, Mendel crossed a plant true-breeding forgreen pods with one true-breeding for yellow pods, as shown inFigure 9-4. The resulting seeds produced an F1 generation that hadonly green-podded plants. No yellow pods developed even thoughone parent had been true-breeding for yellow pods. Only one of thetwo traits found in the P generation appeared in the F1 generation.

Next, Mendel allowed the F1 plants to self-pollinate and plantedthe resulting seeds. When the F2 generation plants grew, heobserved that about three-fourths of the F2 plants had green podsand about one-fourth had yellow pods.

His observations and careful records led Mendel to hypothesizethat something within the pea plants controlled the characteristicsobserved. He called these controls factors. Mendel hypothesizedthat each trait was inherited by means of a separate factor.Because the characteristics studied had two alternative forms, hereasoned that a pair of factors must control each trait.

Recessive and Dominant TraitsWhenever Mendel crossed strains, one of the P traits failed toappear in the F1 plants. In every case, that trait reappeared in aratio of about 3:1 in the F2 generation. This pattern emerged inthousands of crosses and led Mendel to conclude that one factorin a pair may prevent the other from having an effect. Mendelhypothesized that the trait appearing in the F1 generation was con-trolled by a dominant factor because it masked, or dominated, thefactor for the other trait in the pair. He thought that the trait thatdid not appear in the F1 generation but reappeared in the F2 gener-ation was controlled by a recessive factor.

Thus, a trait controlled by a recessive factor had no observableeffect on an organism’s appearance when that trait was paired witha trait controlled by a dominant factor.

The Law of SegregationMendel concluded that the paired factors separate during the for-mation of reproductive cells. That is, each reproductive cell, orgamete, receives one factor of each pair. When two gametes com-bine during fertilization, the offspring have two factors for eachcharacteristic. The law of segregation states that a pair of factorsis segregated, or separated, during the formation of gametes.

The Law of Independent AssortmentMendel also crossed plants that differed in two characteristics,such as flower color and seed color. The data from these more-complex crosses showed that traits produced by dominant factorsdo not necessarily appear together. A green seed pod produced bya dominant factor could appear in a white-flowering pea plant.

Copyright © by Holt, Rinehart and Winston. All rights reserved.

Green-poddedplant

P

F1

F2

3 green-poddedplants

1 yellow-poddedplants

All green-podded plants

Yellow-poddedplant

!

!

34

14

True-breeding green-podded peaplants crossed with true-breedingyellow-podded pea plants produce onlygreen-podded plants. Yet when theF1 generation is permitted to self-pollinate, about one-fourth of the plantsof the F2 generation are yellow-poddedplants.

2

1

FIGURE 9-4

1

2

recessive

from the Latin recessus,meaning “to recede”

Word Roots and Origins

Page 7: CHAPTER 9 FUNDAMENTALS OF GENETICS - … OF GENETICS 177 MENDEL’S RESULTS AND CONCLUSIONS In one of his experiments, Mendel crossed a plant true-breeding for green pods with one

Copyright © by Holt, Rinehart and Winston. All rights reserved.

C H A P T E R 9178

1. Describe what a true-breeding plant is.

2. Outline how Mendel produced plants that had genes for both contrasting traits of acharacteristic.

3. Define the terms dominant and recessive.

4. State in modern terminology the two laws ofheredity that resulted from Mendel‘s work.

5. Differentiate genes from alleles.

CRITICAL THINKING6. Evaluating Results How did Mendel‘s F1 gener-

ation plants differ from his F2 generation plants?

7. Recognizing Relationships Many inherited dis-orders of humans appear in children of parentswho do not have the disorder. How can youexplain this?

8. Applying Information During meiosis, whatallows genes located on the same chromosometo separate independently of one another?

SECTION 1 REVIEW

Mendel concluded that the factors for individual characteristicsare not connected. Recall that the random separation of homolo-gous chromosomes is called independent assortment. The law ofindependent assortment states that factors separate indepen-dently of one another during the formation of gametes.

SUPPORT FOR MENDEL’SCONCLUSIONS

Most of Mendel’s findings agree with what biologists now knowabout molecular genetics. Molecular genetics is the study of thestructure and function of chromosomes and genes. A chromosomeis a threadlike structure made up of DNA. A gene is the segment ofDNA on a chromosome that controls a particular hereditary trait.Because chromosomes occur in pairs, genes also occur in pairs.Each of two or more alternative forms of a gene is called an allele.Mendel’s factors are now called alleles.

Letters are used to represent alleles. Capital letters refer todominant alleles, and lowercase letters refer to recessive alleles.For example, the dominant allele for the trait of purple flower colormay be represented by P, and the recessive allele for the trait ofwhite flower color may be represented by p, as is shown in Figure9-5. Whether a letter is capitalized or lowercased is important. Theactual letter selected to represent an allele is typically the firstletter of the dominant trait. During meiosis, gametes receive onechromosome from each homologous pair of chromosomes. Thus,when the gametes combine in fertilization, the offspring receivesfrom each parent one allele for a given trait.

Mendel’s law of independent assortment is supported by theindependent segregation of chromosomes to gametes during meio-sis. Therefore, the law of independent assortment is observed onlyfor genes located on separate chromosomes or located far apart onthe same chromosome.

P

P Y P y p Y p y

p Y y

Independent assortment of these twopairs of homologous chromosomes (Ppand Yy) would result in gametes thatcontain the allele combinations shownabove. P denotes the dominant purpleflower color, and p denotes the recessivewhite flower color. Y denotes yellow seedcolor, and y denotes green seed color.

FIGURE 9-5

Page 8: CHAPTER 9 FUNDAMENTALS OF GENETICS - … OF GENETICS 177 MENDEL’S RESULTS AND CONCLUSIONS In one of his experiments, Mendel crossed a plant true-breeding for green pods with one

HYPOTHESIS: Genes Can MoveThe prevailing opinion among most geneticists inBarbara McClintock’s time was that genes were linedup on chromosomes in an unchanging way, muchlike beads on a string. However, observations fromher experiments with maize (Zea mays) toldMcClintock otherwise.

As a graduate student at Cornell, McClintockdeveloped new staining techniques and discoveredthat each of the 10 maize chromosomes could be dis-tinguished under the microscope. She noted thatsome changes in the appearance of the corn kernelsand plants were coupled with changes in the shapeof one or more of the maize chromosomes. She alsonoted that kernels that had been exposed to X raysgerminated and grew into seemingly normal plantsbut that the shapes of some of their chromosomeshad changed. She proposed that the maize genomehad a dynamic chromosome repair system thatallowed for growth even after major chromosomedamage that X rays had caused. Geneticists at thetime assumed that a mutated gene was dead andcould not be reactivated. McClintock’s findings,however, challenged this belief.

METHODS: Analyze Maize ChromosomesMcClintock wanted to examine more closely theresults of growing maize that contained broken chro-mosomes. So during the winter of 1944–1945, sheplanted self-pollinated kernels that were the productof many generations of inbreeding and self-fertilization. She hoped to track chromosome repairsby observing changes in chromosome shapes.

RESULTS: Unexpected ChangesWhen the plants germinated, McClintock wasastounded at the results. The leaves had odd patchesthat lacked the normal green coloration. Thesepatches occurred regularly along the leaf blades. She compared chromosomes of these plants withthose of the parent plants under the microscope andconcluded that parts of the offspring plants’ chromo-somes had changed position.

Science in ActionDo Genes Jump?The rediscovery of Mendel’s work in 1900 is said to mark the birthof the science of genetics. Barbara McClintock, born in 1902,devoted her life to this new science. Ironically, certain assump-tions about genetics—which were strongly believed but wrong—prevented early acceptance of McClintock’s exciting conclusions.

179

The colorful maize plant Zeamays was harvested atCornell University duringthe late 1920s and early1930s to study its genet-ics. The varied colors of itskernels worked as a multi-colored spreadsheet ofgenetic data.

R E V I E W

1. What was the purpose ofexposing the maize kernels toX rays?

2. What is a transposon?3. Critical Thinking Why was it

important that McClintockused self-fertilized kernels inher experiments?

www.scilinks.orgTopic: Barbara

McClintockKeyword: HM60924

CONCLUSION: Genes Have theAbility to “Jump”

The changes that McClintock saw inthese plants and their chromosomes led herto conclude that genes are not stable within thechromosome but can move to a new place on achromosome or to a new chromosome entirely.McClintock called these movable genes controllingelements. They were later called transposons.

McClintock observed two kinds of transposons:dissociators and activators. Dissociators could jumpto a new chromosomal location when signaled byactivators. The dissociators would then causechanges in nearby genes on the chromosome and inthe color of the kernels and leaves in the maize.McClintock verified her conclusions through re-peated experiments.

McClintock summarized her findings in 1951 at aCold Spring Harbor symposium, but her work wasnot well received. After many years, recognition ofthe value of her work grew and she was awarded theNobel Prize in medicine or physiology in 1983.

Barbara McClintock

Copyright © by Holt, Rinehart and Winston. All rights reserved.

Page 9: CHAPTER 9 FUNDAMENTALS OF GENETICS - … OF GENETICS 177 MENDEL’S RESULTS AND CONCLUSIONS In one of his experiments, Mendel crossed a plant true-breeding for green pods with one

C H A P T E R 9180

G E N E T I C C R O S S E SToday, geneticists rely on Mendel’s work to predict the likely

outcome of genetic crosses. In this section, you will learn how

to predict the probable genetic makeup and appearance of

offspring resulting from specified crosses.

GENOTYPE AND PHENOTYPEAn organism’s genetic makeup is its genotype (JEEN-uh-TIEP). Thegenotype consists of the alleles that the organism inherits from itsparents. For example, the genotype of the white-flowering peaplant in Figure 9-6 consists of two recessive alleles for white flowercolor, represented as pp. The genotype of a purple-flowering peaplant may be either PP or Pp. Either of these two genotypes wouldresult in a pea plant that has purple flowers because the P allele isdominant.

An organism’s appearance is its phenotype (FEE-noh-TIEP). Thephenotype of a PP or a Pp pea plant is purple flowers, whereas thephenotype of a pp pea plant is white flowers. As this exampleshows, a phenotype does not always indicate genotype. In additionto recessive alleles, certain environmental factors can affect phe-notype. For example, lack of proper nutrition can cause a geneti-cally tall plant to remain short.

SECTION 2

O B J E C T I V E S● Differentiate between the

genotype and the phenotype of an organism.

● Explain how probability is used topredict the results of geneticcrosses.

● Use a Punnett square to predict theresults of monohybrid and dihybridgenetic crosses.

● Explain how a testcross is used toshow the genotype of an individualwhose phenotype expresses thedominant trait.

● Differentiate a monohybrid crossfrom a dihybrid cross.

V O C A B U L A R Ygenotypephenotypehomozygousheterozygous probability monohybrid cross Punnett square genotypic ratio phenotypic ratio testcross complete dominance incomplete dominance codominancedihybrid cross

Copyright © by Holt, Rinehart and Winston. All rights reserved.

The flower color genotype of the peaplant on the left is pp. The plant’sphenotype is white flowers. The flowercolor phenotype of the pea plant on theright is purple flowers. The plant’sgenotype is either Pp or PP.

FIGURE 9-6

Page 10: CHAPTER 9 FUNDAMENTALS OF GENETICS - … OF GENETICS 177 MENDEL’S RESULTS AND CONCLUSIONS In one of his experiments, Mendel crossed a plant true-breeding for green pods with one

181F U N D A M E N T A L S O F G E N E T I C S

When both alleles of a pair are alike, the organism is said to behomozygous (HOH-moh-ZIE-guhs) for that characteristic. An organismmay be homozygous dominant or homozygous recessive. Forexample, a pea plant that is homozygous dominant for flower colorhas the genotype PP. A pea plant that is homozygous recessive forflower color has the genotype pp. When the two alleles in the pairare different, the organism is heterozygous (HET-uhr-OH-ZIE-guhs) forthat characteristic. A pea plant that is heterozygous for flowercolor has the genotype Pp.

PROBABILITYProbability is the likelihood that a specific event will occur. Aprobability may be expressed as a decimal, a percentage, or a frac-tion. Probability is determined by the following equation:

Probability !

For example, in Mendel’s experiments, the dominant trait of yel-low seed color appeared in the F2 generation 6,022 times. Therecessive trait of green seed color appeared 2,001 times. The totalnumber of individuals was 8,023 (6,022 " 2,001). Using the prob-ability equation above we can determine that the probability thatthe dominant trait will appear in a similar cross is

#68,,002223

# ! 0.75

Expressed as a percentage, the probability is 75 percent. Expressedas a fraction, the probability is 3/4.

The probability that the recessive trait will appear in an F2generation is

#28,,000213

# ! 0.25

Expressed as a percentage, the probability is 25 percent. Expressedas a fraction, the probability is 1/4. Fractions can also be expressedas ratios. For example, the ratio 1:3 represents the same probabil-ity that 1/4 does. Probability tells us that there are three chancesin four that an offspring of two heterozygous individuals will havethe dominant trait and one chance in four that the offspring willhave only the recessive trait.

The results predicted by probability are more likely to occurwhen there are many trials. For example, many coin tosses shouldyield a result of heads 50 percent of the time and tails 50 percentof the time. However, if you toss a coin only a few times, you mightnot get this result. But each time a coin is tossed, the probability oflanding tails is 50 percent. Only after many, many tries would yoube likely to get the percentage of heads predicted on the basis ofprobability, that is, 50 percent heads and 50 percent tails.

number of times an event is expected to happen######

number of times an event could happen

Copyright © by Holt, Rinehart and Winston. All rights reserved.

Calculating ProbabilityMaterials paper sack containing20 jelly beans of three different col-ors (with an unknown number ofeach color)

Procedure1. Obtain a sack of 20 jelly beans

from your teacher. Do not lookinto the sack. Do not eat thejelly beans. There are three pos-sible colors of jelly beans thatcan be pulled from the sack. Pullone jelly bean out, and recordthe color. Return the jelly beanto the sack, and shake the bagto mix the jelly beans.

2. Repeat step 1 until you haveexamined 20 jelly beans.

3. Determine the probability of getting a jelly bean of a specificcolor with a single draw. Do thisfor each of the three colors ofjelly beans. Compare your resultswith those of the rest of theclass.

Analysis Does anyone have thesame probabilities that you do? Are any probabilities very close toyours? Are any probabilities very dif-ferent from yours? From these obser-vations, determine how many jellybeans of each color are in your sack.

Quick Lab

Page 11: CHAPTER 9 FUNDAMENTALS OF GENETICS - … OF GENETICS 177 MENDEL’S RESULTS AND CONCLUSIONS In one of his experiments, Mendel crossed a plant true-breeding for green pods with one

C H A P T E R 9182

PREDICTING RESULTS OFMONOHYBRID CROSSES

A cross in which only one characteristic is tracked is amonohybrid (MAHN-oh-HIE-brid) cross. The offspring of amonohybrid cross are called monohybrids. A cross betweena pea plant that is true-breeding for producing purple flow-ers and one that is true-breeding for producing white flow-ers is an example of a monohybrid cross. Biologists use adiagram called a Punnett (PUHN-uht) square, such as the oneshown in Figure 9-7, to aid them in predicting the probabledistribution of inherited traits in the offspring. The followingexamples show how a Punnett square can be used to predictthe outcome of different types of crosses.

Example 1: Homozygous ! HomozygousFigure 9-7 shows a cross between a pea plant homozygous forpurple flower color (PP) and a pea plant homozygous for whiteflower color (pp). The alleles carried in gametes of the homozy-gous dominant parent are represented by P’s on the left side of thePunnett square. The alleles carried in gametes of the homozygousrecessive parent are represented by p’s across the top of thePunnett square. Each box within the Punnett square is filled in withthe letters, or alleles, that are above it and at left of it outside thesquare. The combinations of alleles in the four boxes indicate thepossible genotypes that can result from the cross. The predictedgenotype is Pp in every case. Thus, there is a 100 percent proba-bility that the offspring will have the genotype Pp and thus the phe-notype purple flower color.

Example 2: Homozygous ! HeterozygousFigure 9-8 shows a cross between a guinea pig that is homozygousdominant for the trait of black coat color (BB) and a guinea pig thatis heterozygous for this trait (Bb). The letter b stands for the reces-

sive allele. Genotype bb results in a brown coat. Notice thattwo possible genotypes can result from this cross: BB or Bb.The probability of an offspring having the genotype BB is2/4, or 50 percent. The probability of an offspring having thegenotype Bb is also 2/4, or 50 percent. You could expectabout 50 percent of the offspring resulting from this cross tobe homozygous dominant for the black coat and about 50percent to be heterozygous dominant for a black coat. Theprobable phenotype is black coat color in every case; thus,4/4, or 100 percent, of the offspring are expected to have ablack coat. What would happen if the homozygous guineapig were homozygous recessive for coat color? The homozy-gote would have the genotype bb. Crossing a bb guinea pigwith a Bb guinea pig is likely to produce about 50 percent Bboffspring and about 50 percent bb offspring.

B b

B

(BB)

(Bb)

B

BB Bb

BB Bb

p p

P

(PP)

(pp)

P

Pp Pp

Pp Pp

A pea plant homozygous for purpleflowers that is crossed with a pea planthomozygous for white flowers willproduce only purple-flowering offspring.Note that all of the offspring, calledmonohybrids, are heterozygous forflower color.

FIGURE 9-7

Crossing a guinea pig homozygous forblack coat color with one heterozygousfor black coat color produces all black-coated monohybrid offspring. Note thathalf of the monohybrid offspring arepredicted to be homozygous for coatcolor.

FIGURE 9-8

Copyright © by Holt, Rinehart and Winston. All rights reserved.

Page 12: CHAPTER 9 FUNDAMENTALS OF GENETICS - … OF GENETICS 177 MENDEL’S RESULTS AND CONCLUSIONS In one of his experiments, Mendel crossed a plant true-breeding for green pods with one

183F U N D A M E N T A L S O F G E N E T I C S

The probable results of crossing tworabbits that are heterozygous for blackcoat color are 50 percent heterozygousblack individuals, 25 percenthomozygous black individuals, and 25 percent homozygous brownindividuals.

FIGURE 9-9

B b

B

(Bb)

(Bb)

b

BB Bb

Bb bb

b b

B

B

Bb Bb

Bb Bb

b b

B

b

Bb Bb

bb bb

If a black guinea pig is crossed with abrown guinea pig and even one of theoffspring is brown, the black guinea pigis heterozygous for coat color.

FIGURE 9-10

Example 3: Heterozygous ! HeterozygousIn rabbits, the allele for black coat color (B) is dominant over theallele for brown coat color (b). The Punnett square in Figure 9-9shows the predicted results of crossing two rabbits that are het-erozygous (Bb) for coat color. As you can see, 1/4 (25 percent) ofthe offspring are predicted to have the genotype BB, 1/2 (50 per-cent) are predicted to have the genotype Bb, and 1/4 (25 percent)are predicted to have the genotype bb. Thus, 3/4 (75 percent) ofthe offspring resulting from this cross are predicted to have a blackcoat. One-fourth (25 percent) of the offspring are predicted to havea brown coat.

The ratio of the genotypes that appear in offspring is called thegenotypic ratio. The probable genotypic ratio of the monohybridcross represented in Figure 9-9 is 1 BB : 2 Bb : 1 bb. The ratio of theoffspring’s phenotypes is called the phenotypic ratio. The prob-able phenotypic ratio of the cross represented in Figure 9-9 is 3 black : 1 brown.

Example 4: TestcrossRecall that in guinea pigs, both BB and Bb result in a black coat.How might you determine whether a black guinea pig is homozy-gous (BB) or heterozygous (Bb)? You could perform a testcross, inwhich an individual of unknown genotype is crossed with ahomozygous recessive individual. A testcross can determine thegenotype of any individual whose phenotype expresses the domi-nant trait. You can see from Figure 9-10 that if the black guinea pigof unknown genotype is homozygous black, all offspring will beblack. If the individual with the unknown genotype is heterozygousblack, about half of the offspring will be black. In reality, if the crossproduced any brown offspring, the genotype of the black-coatedparent is likely to be heterozygous.

Copyright © by Holt, Rinehart and Winston. All rights reserved.

Page 13: CHAPTER 9 FUNDAMENTALS OF GENETICS - … OF GENETICS 177 MENDEL’S RESULTS AND CONCLUSIONS In one of his experiments, Mendel crossed a plant true-breeding for green pods with one

C H A P T E R 9184

R r

R

(Rr)

(Rr)

r

RR Rr

Rr rr

When red-flowering four o’clocks arecrossed with white-flowering fouro’clocks, all of the F1 offspring producepink flowers, an intermediate betweenthe two parental phenotypes. When the F1 generation is interbred, red-flowering, pink-flowering, and white-flowering plants are produced becausethe trait for red flower color hasincomplete dominance over the trait for white flower color.

FIGURE 9-11

www.scilinks.orgTopic: DominanceKeyword: HM60422

Example 5: Incomplete DominanceRecall that in Mendel’s pea-plant crosses, one allele was completelydominant over another, a relationship called complete dominance.In complete dominance, heterozygous plants and homozygousdominant plants are indistinguishable in phenotype. For example,both pea plants PP and Pp for flower color have purple flowers.

Sometimes, however, the F1 offspring will have a phenotype inbetween that of the parents, a relationship called incompletedominance. Incomplete dominance occurs when the phenotype ofa heterozygote is intermediate between the phenotypes deter-mined by the dominant and recessive traits. In four o’clocks, forexample, both the allele for red flowers (R) and the allele for whiteflowers (r) influence the phenotype. Neither allele is completelydominant over the other allele. When four o’clocks self-pollinate, red-flowering plants produce only red-flowering offspringand white-flowering plants produce only white-flowering offspring.However, when red four o’clocks are crossed with white fouro’clocks, all of the F1 offspring have pink flowers. One hundred per-cent of the offspring of this cross have the Rr genotype, whichresults in a pink phenotype.

What would be the result of crossing two pink-flowering (Rr) fouro’clocks? As the Punnett square in Figure 9-11 shows, the probablegenotypic ratio is 1 RR : 2 Rr : 1 rr. Given that neither the allele forred flowers (R) nor the allele for white flowers (r) is completelydominant, the probable phenotypic ratio is 1 red : 2 pink : 1 white.

Example 6: Codominance Codominance occurs when both alleles for a gene are expressed ina heterozygous offspring. In codominance, neither allele is domi-nant or recessive, nor do the alleles blend in the phenotype.

For example, the three human MN blood types, M, N, and MN,are determned by two alleles, LM, and LN. The letters M and N referto two molecules on the surface of the red blood cell. The genotypeof a person with blood type MN is LMLN, and neither allele is domi-nant over the other. Type MN blood cells cary both M- and N-typesof molecules on their surface.

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185F U N D A M E N T A L S O F G E N E T I C S

Determining GenotypesMaterials pencil and paperProcedure The ability to roll thetongue upward from the sides is adominant, inherited trait. In onefamily, both parents and three chil-dren are tongue rollers, while onechild is not. Determine the genotypeand phenotype of each parent.Analysis Are the parents homozygous or heterozygous? Are the children homozygous orheterozygous?

Quick Lab

ry ry ry ry

RY

RY

RY

RY

RrYy RrYy RrYy RrYy

RrYy RrYy RrYy RrYy

RrYy RrYy RrYy RrYy

RrYy RrYy RrYy RrYy

RRYY

rryy

This Punnett square shows a dihybridcross between a pea plant that ishomozygous recessive for wrinkled,green seeds (rryy ) and a pea plant thatis homozygous dominant for round,yellow seeds (RRYY ).

FIGURE 9-12

PREDICTING RESULTS OFDIHYBRID CROSSES

A dihybrid (die-HIE-brid) cross is a cross in which two characteristicsare tracked. The offspring of a dihybrid cross are called dihybrids.Predicting the results of a dihybrid cross is more complicated thanpredicting the results of a monohybrid cross because more combi-nations of alleles are possible. For example, to predict the results ofa cross in which both seed texture and seed color are tracked, youhave to consider how four alleles from each parent can combine.

Homozygous ! HomozygousSuppose that you want to predict the results of a cross between apea plant that is homozygous for round, yellow seeds and one thatis homozygous for wrinkled, green seeds. In pea plants, the allelefor round seeds (R) is dominant over the allele for wrinkled seeds(r), and the allele for yellow seeds (Y ) is dominant over the allelefor green seeds (y).

As Figure 9-12 shows, the Punnett square used to predict theresults of a cross between a parent of the genotype RRYY and a par-ent of the genotype rryy will contain 16 boxes. Alleles are carriedby the male and female gametes (pollen and ovule). The indepen-dently sorted alleles from one parent—RY, RY, RY, and RY—arelisted along the left side of the Punnett square. The independentlysorted alleles from the other parent—ry, ry, ry, and ry—are listedalong the top of the Punnett square. Each box is filled with the let-ters that are above it and to the left of it outside the square. Noticethat the genotype of all of the offspring of this cross will be het-erozygous for both traits (RrYy); therefore, all of the offspring willhave round, yellow seed phenotypes.

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C H A P T E R 9186

Heterozygous ! HeterozygousThe same procedure is used to determine the results of crossingtwo pea plants heterozygous for round, yellow seeds. As Figure 9-13 shows, the offspring of this dihybrid cross are likely to havenine different genotypes. These nine genotypes will result in peaplants that have the following four phenotypes:• 9/16 that have round, yellow seeds (genotypes RRYY, RRYy, RrYY,

and RrYy)• 3/16 that have round, green seeds (genotypes RRyy and Rryy)• 3/16 that have wrinkled, yellow seeds (genotypes rrYY and rrYy)• 1/16 that have wrinkled, green seeds (genotype rryy)

1. Explain why a phenotype might not always indi-cate genotype.

2. Identify the equation used to determineprobability.

3. Explain how you might go about determiningthe genotype of a purple-flowering plant.

4. Illustrate in the form of a Punnett square theresults of crossing a pink-flowering four o’clockwith a white-flowering four o’clock.

5. Explain the difference between a monohybridcross and a dihybrid cross and give an exampleof each.

CRITICAL THINKING6. Analyzing Concepts The offspring of two short-

tailed cats have a 25 percent chance of havingno tail, a 25 percent chance of having a longtail, and a 50 percent chance of having a shorttail. Using this information, what can youhypothesize about the genotypes of the parentsand the way in which tail length is inherited?

7. Inferring Relationships If you crossed twopurple-flowering pea plants and all of the F1 off-spring were purple-flowering, what could you sayabout the genotypes of the parents? If some ofthe F1 offspring were white-flowering, what couldyou say about the genotypes of the parents?

SECTION 2 REVIEW

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www.scilinks.orgTopic: Punnett SquaresKeyword: HM61249

RY Ry

RrYy

RrYy

rY ry

RY

Ry

rY

ry

RRYY RRYy RrYY RrYy

RRYy RRyy RrYy Rryy

RrYY RrYy rrYY rrYy

RrYy Rryy rrYy rryy

A dihybrid cross of two individualsheterozygous for both traits of interestis likely to result in nine differentgenotypes and four differentphenotypes.

FIGURE 9-13

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CHAPTER HIGHLIGHTS

187F U N D A M E N T A L S O F G E N E T I C S

genetics (p. 173)heredity (p. 173)trait (p. 173)pollination (p. 174)self-pollination (p. 174)

cross-pollination (p. 174)true-breeding (p. 175)P generation (p. 175)F1 generation (p. 175)F2 generation (p. 175)

dominant (p. 177)recessive (p. 177)law of segregation (p. 177)law of independent

assortment (p. 178)

molecular genetics (p. 178)allele (p. 178)

Vocabulary

Mendel’s LegacySECTION 1

● The study of how characteristics are transmitted fromparents to offspring is called genetics.

● Mendel observed seven characteristics of pea plants.Each characteristic occurred in two contrasting traits.

● Self-pollination, in which pollen is transferred from theanthers of a flower to either the stigma of the sameflower or the stigma of another flower on the sameplant, normally occurs in pea plants. Cross-pollinationoccurs when pollen is transferred between flowers of two different plants.

● Mendel concluded that inherited characteristics arecontrolled by factors that occur in pairs. In hisexperiments on pea plants, one factor in a pair maskedthe other. The trait that masked the other was called thedominant trait. The trait that was masked was called therecessive trait.

● The law of segregation states that a pair of factors issegregated, or separated, during the formation ofgametes. Two factors for a characteristic are thencombined when fertilization occurs and a new offspring is produced.

● The law of independent assortment states that factors forindividual characteristics are distributed to gametesindependently. The law of independent assortment isobserved only for genes that are located on separatechromosomes or are far apart on the same chromosome.

● We now know that the factors that Mendel studied arealleles, or alternative forms of a gene. Each of two ormore alternative forms of a gene is called an allele. Oneallele for each trait is passed from each parent to theoffspring.

Genetic CrossesSECTION 2

genotype (p. 180)phenotype (p. 180)homozygous (p. 181)heterozygous (p.181)

probability (p. 181)monohybrid cross (p. 182)Punnett square (p. 182)genotypic ratio (p. 183)

phenotypic ratio (p. 183)testcross (p. 183)complete dominance

(p. 184)

incomplete dominance (p. 184)

codominance (p. 184)dihybrid cross (p. 185)

Vocabulary

● The genotype is the genetic makeup of an organism. Thephenotype is the appearance of an organism.

● Probability is the likelihood that a specific event willoccur. A probability may be expressed as a decimal, apercentage, or a fraction.

● A Punnett square can be used to predict the outcome ofgenetic crosses.

● A cross in which one characteristic is tracked is amonohybrid cross. The offspring of a monohybrid crossare called monohybrids.

● A testcross, in which an individual of unknown genotypeis crossed with a homozygous recessive individual, can beused to determine the genotype of an individual whosephenotype expresses the dominant trait.

● Complete dominance occurs when heterozygousindividuals and dominant homozygous individuals areindistinguishable in phenotype.

● Incomplete dominance occurs when two or more allelesinfluence the phenotype and results in a phenotypeintermediate between the dominant trait and therecessive trait.

● Codominance occurs when both alleles for a gene areexpressed in a heterozygous offspring. Neither allele isdominant or recessive, nor do the alleles blend in thephenotype as they do in incomplete dominance.

● A cross in which two characteristics are tracked is adihybrid cross. The offspring of a dihybrid cross are called dihybrids.

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CHAPTER REVIEW

C H A P T E R 9188

USING VOCABULARY1. For each pair of terms, explain how the meanings

of the terms differ.a. homozygous and heterozygousb. law of segregation and law of independent

assortmentc. genetics and heredity

2. Use the following terms in the same sentence: pollination, self-pollination, andcross-pollination.

3. For each pair of terms, explain the relationshipbetween the terms.a. genotype and phenotypeb. monohybrid cross and dihybrid crossc. allele and trait

4. Word Roots and Origins The word dominantcomes from the Latin dominari, which means “to rule.” Using this information, explain why theterm dominant is a good name for the geneticphenomenon that the term describes.

UNDERSTANDING KEY CONCEPTS5. Relate why Mendel began his experiments by

allowing pea plants to self-pollinate for severalgenerations.

6. Explain the difference between dominant andrecessive traits.

7. Describe the differences between the P genera-tion, the F1 generation, and the F2 generation.

8. Relate why it is not necessary, when the domi-nant and recessive traits are known, to use theterm homozygous when referring to the genotypeof an individual that has a recessive phenotype.

9. State the difference between a monohybrid crossand a dihybrid cross.

10. Propose how crossing-over during meiosis mightaffect the segregation of genes that are on thesame chromosome.

11. Relate the events of meiosis to the law of segregation.

12. Summarize how a gardener who has a pea plantthat produces round seeds can determinewhether the plant is homozygous or heterozy-gous for the allele that determines seed texture.(In pea plants, round seed texture is dominantover wrinkled seed texture.)

13. Relate probability to the study of genetics.

14. Predict the results of a cross between a rabbithomozygous dominant for black coat color (BB)and a rabbit homozygous recessive for browncoat color (bb).

15. CONCEPT MAPPING Use the followingterms to create a concept map that

illustrates information related to Mendel’sexperiments: pea plants, heredity, self-pollination, cross-pollination, F1 generation,F2 generation, trait, and true-breeding.

CRITICAL THINKING16. Interpreting Graphics Use the Punnett square

below to answer the questions that follow.a. Does the Punnett square demonstrate a

monohybrid cross or a dihybrid cross?b. List the genotypes of the parents.c. Give the genotypic ratio predicted by the

Punnett square for the cross.

17. Predicting Results In rabbits, the allele for blackcoat color (B) is dominant over the allele forbrown coat color (b). Predict the results of across between a rabbit homozygous for blackcoat color (BB) and a rabbit homozygous forbrown coat color (bb).

18. Applying Information Write areport summarizing how an under-standing of heredity allows animal

breeders to develop animals that have desirabletraits. Find out what kinds of animals are bred forspecial purposes.

QT Qt qT qt

QT

Qt

qT

qt

QQTT QQTt QqTT QqTt

QQTt QQtt QqTt Qqtt

QqTT QqTt qqTT qqTt

QqTt Qqtt qqTt qqtt

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189F U N D A M E N T A L S O F G E N E T I C S

Standardized Test PreparationDIRECTIONS: Choose the letter of the answer choicethat best answers the question.

1. What is a procedure in which an individual ofunknown genotype is crossed with a homozygousrecessive individual to determine the genotype ofthe unknown individual called?A. a monohybrid crossB. a dihybrid crossC. a hybrid crossD. a testcross

2. In a monohybrid cross of two heterozygous par-ents (Pp), what would the expected genotypes ofthe offspring be?F. 1 PP : 2 Pp : 1 ppG. 1 pp : 3 PPH. 3 Pp : 1 ppJ. all Pp

3. Which of the following is an example of a geno-type of a heterozygous individual? A. pB. YYC. ZzD. rr

INTERPRETING GRAPHICS: Use the diagrams ofchromosomes below to answer the question that fol-lows. The single chromosome below has two genes,both of which carry a dominant allele Q and R.

4. Homologous chromosomes are chromosomesthat carry genes for the same characteristics,such as eye color or hair color. Which of thechromosomes in the bottom row could not bethe homologous chromosome for the single chromosome in the top row?F. 1G. 2H. 3J. 4

DIRECTIONS: Complete the analogy below:5. Rr : genotype :: red :

A. F1 generationB. heterozygoteC. phenotypeD. dominant

INTERPRETING GRAPHICS: Use the diagram of aPunnett square below to answer the question thatfollows.

The Punnett square above shows the expectedresults of a cross between two pea plants. R and rrepresent the alleles for round seed and wrinkledseed traits, respectively.

6. What would the seed texture phenotype of theplant in box 4 be?F. roundG. RrH. wrinkledJ. rr

SHORT RESPONSEMendel was able to observe certain traits as theywere passed on by carefully controlling how the peaplants were pollinated. Explain why Mendel began hisexperiments by allowing pea plants to self-pollinatefor several generations.

EXTENDED RESPONSEA cross between two pea plants that have axial flow-ers and inflated pods gives the following offspring: 20 that have axial flowers and inflated pods, 7 withaxial flowers and constricted pods, and 5 that haveterminal flowers and inflated pods.

Part A Identify the most probable genotype of thetwo parents.

Part B Use a Punnett square to explain the results.

Q

R

1 2 3 4Q

R

q

R

q

q

q

r

r r

R

r

1 2

3 4

Before answering word problemsthat are genetics questions, write the problem downby using letters to symbolize genotypes.

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C H A P T E R 9190

Modeling Monohybrid Crosses

■ Predict the genotypic and phenotypic ratios of off-spring resulting from the random pairing of gametes.

■ Calculate the genotypic ratio and phenotypic ratioamong the offspring of a monohybrid cross.

■ predicting■ organizing■ analyzing data■ calculating

■ lentils■ green peas■ 2 Petri dishes

Background1. How many traits are involved in a monohybrid

cross? How many alleles are involved? 2. What prevents the expression of a recessive allele?3. When gametes form, what happens to the alleles

for each trait?

Simulating a Monohybrid Cross

1. You will model the random pairing of alleles bychoosing lentils and peas from Petri dishes. Thesedried seeds will represent the alleles for seed color.A green pea will represent G, the dominant allelefor green seeds, and a lentil will represent g, therecessive allele for yellow seeds.

2. The seeds in each Petri dish will represent thealleles from a single parent. Label one Petri dish“female gametes” and the other Petri dish “malegametes.” Place one green pea and one lentil in thePetri dish labeled “female gametes,” and place onegreen pea and one lentil in the Petri dish labeled“male gametes.”

3. Each parent contributes one allele to each offspring.Model a cross between these two parents by choos-ing a random pairing of the dried seeds from thetwo containers. Do so by simultaneously picking oneseed from each container without looking. Place thepair of seeds together on the lab table. The pair ofseeds represents the genotype of one offspring.

4. Record the genotype of the first offspring in yourlab report in a table like Table A, shown below.

5. Return the seeds to their original dishes, and repeatstep 3 nine more times. Record the genotype ofeach offspring in your data table.

6. Based on each offspring’s genotype, determine andrecord each offspring’s phenotype. Assume that theallele for green seeds, G, is completely dominantover the allele for yellow seeds, g.

PART A

MATERIALS

PROCESS SKILLS

OBJECTIVES

EXPLORATION LAB

TABLE A GAMETE PAIRINGS

Trial Offspring genotype Offspring phenotype

1

2

3

4

5

6

7

8

9

10

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191F U N D A M E N T A L S O F G E N E T I C S

Calculating Genotypic andPhenotypic Ratios

7. In your lab report, prepare a data table similar to Table B, shown below.

8. Determine the genotypic and phenotypic ratios among the offspring. First, count and record the number of homozygous dominant, heterozygous,and homozygous recessive individuals recorded inTable A. Then, record the number of offspring that produce green seeds and the number that produce yellow seeds under “Phenotypes” in your data table.

9. Calculate the genotypic ratio for each genotype byusing the following equation:

Genotypic ratio !

10. Calculate the phenotypic ratio for each phenotype byusing the following equation:

Phenotypic ratio !

11. Now, pool the data for the whole class, and record the data in your lab report in a table like Table C.

12. Compare your class’s sample with your small sampleof 10. Calculate and record in your data table thegenotypic and phenotypic ratios for the class data.

13. Construct a Punnett square showing the parents and their offspring in your lab report.

14. Clean up your materials before leaving the lab.

Analysis and Conclusions 1. What characteristic is being studied in this investi-

gation?2. What are the genotypes of the parents? Describe

the genotypes of both parents by using the termshomozygous, heterozygous, or both.

3. What does each seed in the Petri dish represent? 4. When the seeds were selected and paired, what did

the pairs represent? 5. Did Tables B and C reflect a classic monohybrid-

cross phenotypic ratio of 3:1? 6. When the class data were tabulated, did a classic

monohybrid-cross phenotypic ratio of 3:1 result? 7. If a genotypic ratio of 1:2:1 is observed, what must

the genotypes of both parents be? 8. Show what the genotypes of the parents would

be if 50 percent of the offspring were green and 50 percent of the offspring were yellow.

9. Construct a Punnett square for the cross of a heterozygous black guinea pig and an unknownguinea pig whose offspring include a recessive white-furred individual. What are the possible geno-types of the unknown parent?

Further InquiryDesign a model to demonstrate a dihybrid cross of twoparents that are heterozygous for two characteristics.Construct and complete a Punnett square for this cross.

number of offspring with a given phenotype""""

total number of offspring

number of offspring with a given genotype""""

total number of offspring

PART B

TABLE C OFFSPRING RATIOS (Entire Class)

Genotypes Total Genotypic ratio

Homozygousdominant (GG )

Heterozygous(Gg) _____ : _____ : _____Homozygousrecessive (gg)

Phenotypes Phenotypic ratio

Greenseeds _____ : _____Yellow seeds

TABLE B OFFSPRING RATIOS

Genotypes Total Genotypic ratio

Homozygousdominant (GG )

Heterozygous(Gg) _____ : _____ : _____Homozygousrecessive (gg)

Phenotypes Phenotypic ratio

Greenseeds _____ : _____ Yellow seeds

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