Unit 4 Chapter 15

Chapter
15
The Chromosomal Basis of
Inheritance
PowerPoint Lecture Presentations for
Biology
Eighth Edition
Neil Campbell and Jane Reece
Lectures by Chris Romero, updated by Erin Barley with contributions from Joan Sharp
Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Overview: Locating Genes Along

Chromosomes
Mendels hereditary factors were genes,
though this wasnt known at the time
Today we can show that genes are located on
chromosomes
The location of a particular gene can be seen
by tagging isolated chromosomes with a
fluorescent dye that highlights the gene
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings
Fig. 15-1
Concept 15.1: Mendelian inheritance has its

physical basis in the behavior of chromosomes
Mitosis and meiosis were first described in the
late 1800s
The chromosome theory of inheritance
states:
Mendelian genes have specific loci (positions) on
chromosomes
Chromosomes undergo segregation and independent
assortment
The behavior of chromosomes during meiosis

was said to account for Mendels laws of
segregation and independent assortment
Fig. 15-2
P Generation
Yellow-round
seeds (YYRR)
Y
R
Green-wrinkled
seeds (yyrr)
y
r
Meiosis
Fertilization
R Y
Gametes
All F1 plants produce

yellow-round seeds (YyRr)
F1 Generation
LAW OF SEGREGATION
The two alleles for each gene
separate during gamete
formation.
LAW OF INDEPENDENT
ASSORTMENT Alleles of genes
on nonhomologous
chromosomes assort
independently during gamete
formation.
Meiosis
R
Metaphase I
1
R
Anaphase I
Metaphase II
Gametes
R
1
F2 Generation
/4
YR
y
r
r
1
/4
r
1
yr
/4
Yr
y
R
R
1
/4
yR
An F1 F1 cross-fertilization
:3
:3
:1
Fig. 15-2a
Green-wrinkled
seeds (yyrr)
Yellow-round
seeds (YYRR)
P Generation
Y
Y
R R
y
r
Meiosis
Gametes
R Y
Fertilization

Fig. 15-2b

0.5 mm
F1 Generation
LAW OF SEGREGATION
The two alleles for each gene
separate during gamete
formation.
r
Y
LAW OF INDEPENDENT
ASSORTMENT Alleles of genes
on nonhomologous
chromosomes assort
independently during gamete
formation.
Meiosis
r
Metaphase I
Y
1
r
Anaphase I
Y
Metaphase II
Gametes
Y
R
YR
yr
Yr
y
R
R
1
yR
3
Fig. 15-2c
F2 Generation
An F1 F1 cross-fertilization
3
9
:3
:3
:1
Morgans Experimental Evidence:

Scientific Inquiry
The first solid evidence associating a specific
gene with a specific chromosome came from
Thomas Hunt Morgan, an embryologist
Morgans experiments with fruit flies provided
convincing evidence that chromosomes are the
location of Mendels heritable factors
Morgans Choice of Experimental

Organism
Several characteristics make fruit flies a
convenient organism for genetic studies:
They breed at a high rate
A generation can be bred every two weeks
They have only four pairs of chromosomes
Morgan noted wild type, or normal,

phenotypes that were common in the fly
populations
Traits alternative to the wild type are called
mutant phenotypes
Fig. 15-3
Correlating Behavior of a Genes Alleles with

Behavior of a Chromosome Pair
In one experiment, Morgan mated male flies
with white eyes (mutant) with female flies with
red eyes (wild type)
The F1 generation all had red eyes
The F2 generation showed the 3:1 red:white
eye ratio, but only males had white eyes
Morgan determined that the white-eyed mutant

allele must be located on the X chromosome
Morgans finding supported the chromosome
theory of inheritance
Fig. 15-4
EXPERIMENT
P
Generation
F1
Generation
All offspring
had red eyes
RESULTS
F2
Generation
CONCLUSION
P
Generation
w+
X
X
w+
X
Y
w
Eggs
F1
Generation
w+
Sperm
w+
w+
w
w+
Eggs
F2
Generation
w+
w
Sperm
w+
w+
w+
w
w+
Fig. 15-4a
EXPERIMENT
P
Generation
F1
Generation
All offspring
had red eyes
Fig. 15-4b
RESULTS
F2
Generation
Fig. 15-4c
CONCLUSION
P
Generation
w+
X
X
w+
X
Y
Eggs
F1
Generation
Sperm
w+
w+
w+
w
w+
Eggs
F2
Generation
w+
Sperm
w+
w
w
w+
w+
Concept 15.2: Sex-linked genes exhibit

unique patterns of inheritance
In humans and some other animals, there is a
chromosomal basis of sex determination
The Chromosomal Basis of

Sex
In humans and other mammals, there are two
varieties of sex chromosomes: a larger X
chromosome and a smaller Y chromosome
Only the ends of the Y chromosome have
regions that are homologous with the X
chromosome
The SRY gene on the Y chromosome codes for
the development of testes
Fig. 15-5
Females are XX, and males are XY

Each ovum contains an X chromosome, while
a sperm may contain either an X or a Y
chromosome
Other animals have different methods of sex
determination
Fig. 15-6
44 +
XY
44 +
XX
Parents
22 +
22 +
or Y
X
Sperm
44 +
XX
or
22 +
X
Egg
44 +
XY
Zygotes (offspring)
(a) The X-Y system
22 +
XX
22 +
X
76 +
ZW
76 +
ZZ
32
(Diploid)
16
(Haploid)
(b) The X-0 system
(c) The Z-W system
(d) The haplo-diploid system
Fig. 15-6a
44 +
XY
44 +
XX
Parents
22 +
22 +
or
X
Y
Sperm
44 +
XX
22 +
X
Egg
or
44 +
XY
Zygotes (offspring)
(a) The X-Y system
Fig. 15-6b
22 +
XX
(b) The X-0 system
22 +
X
Fig. 15-6c
76 +
ZW
(c) The Z-W system
76 +
ZZ
Fig. 15-6d
32
(Diploid)
(d) The haplo-diploid system
16
(Haploid)
Inheritance of Sex-Linked
Genes
The sex chromosomes have genes for many
characters unrelated to sex
A gene located on either sex chromosome is
called a sex-linked gene
In humans, sex-linked usually refers to a gene
on the larger X chromosome
Sex-linked genes follow specific patterns of

inheritance
For a recessive sex-linked trait to be expressed
A female needs two copies of the allele
A male needs only one copy of the allele
Sex-linked recessive disorders are much more

common in males than in females
Fig. 15-7
XNXN
Sperm Xn
X nY
Sperm XN
Eggs XN
XNXn XNY
XN
XNXn XNY
(a)
XNXn
XNY
XNXn
Sperm Xn
X nY
Eggs XN XNXN XNY
Eggs XN
XNXn XNY
XnXN XnY
Xn
X nX n X nY
Xn
(b)
(c)
Some disorders caused by recessive alleles on

the X chromosome in humans:
Color blindness
Duchenne muscular dystrophy
Hemophilia
X Inactivation in Female
Mammals
In mammalian females, one of the two X
chromosomes in each cell is randomly
inactivated during embryonic development
The inactive X condenses into a Barr body
If a female is heterozygous for a particular
gene located on the X chromosome, she will be
a mosaic for that character
Fig. 15-8
X chromosomes
Early embryo:
Two cell
populations
in adult cat:
Active X
Allele for
orange fur
Allele for
black fur
Cell division and

X chromosome
inactivation
Active X
Inactive X
Black fur
Orange fur
Concept 15.3: Linked genes tend to be inherited

together because they are located near each other on
the same chromosome
Each chromosome has hundreds or thousands
of genes
Genes located on the same chromosome that
tend to be inherited together are called linked
genes
How Linkage Affects

Inheritance
Morgan did other experiments with fruit flies to
see how linkage affects inheritance of two
characters
Morgan crossed flies that differed in traits of
body color and wing size
Fig. 15-UN1
b+vg+
Parents
in testcross
Most
offspring
bvg
bvg
bvg
b+vg+
bvg
or
bvg
bvg
Fig. 15-9-1
EXPERIMENT
P Generation (homozygous)
Wild type
(gray body,
normal wings)
b+b+vg+vg+
Double mutant
(black body,
vestigial wings)
bbvgvg
Fig. 15-9-2
EXPERIMENT
Wild type
(gray body,
normal wings)
bbvgvg
b+b+vg+vg+
F1 dihybrid
(wild type)
b+bvg+vg
Double mutant
(black body,
vestigial wings)
TESTCROSS
Double mutant
bbvgvg
Fig. 15-9-3
EXPERIMENT
Wild type
(gray body,
normal wings)
Double mutant
(black body,
vestigial wings)
bbvgvg
b+b+vg+vg+
F1 dihybrid
(wild type)
Double mutant
TESTCROSS
bbvgvg
b+bvg+vg
Testcross
offspring
b+vg+
bvg
Wild type
(gray-normal)
Blackvestigial
Eggs
b+vg
bvg+
Grayvestigial
Blacknormal
bvg
Sperm
b+bvg+vg bbvgvg b+bvgvg bbvg+vg
Fig. 15-9-4
EXPERIMENT
Wild type
(gray body,
normal wings)
Double mutant
(black body,
vestigial wings)
bbvgvg
b+b+vg+vg+
F1 dihybrid
(wild type)
Double mutant
TESTCROSS
bbvgvg
b+bvg+vg
Testcross
offspring
b+vg+
bvg
Wild type
(gray-normal)
Blackvestigial
Eggs
b+vg
bvg+
Grayvestigial
Blacknormal
bvg
Sperm
b+bvg+vg bbvgvg b+bvgvg bbvg+vg

PREDICTED RATIOS
If genes are located on different chromosomes:
If genes are located on the same chromosome and

parental alleles are always inherited together:
965
944
206
185
RESULTS
Morgan found that body color and wing size

are usually inherited together in specific
combinations (parental phenotypes)
He noted that these genes do not assort
independently, and reasoned that they were on
the same chromosome
However, nonparental phenotypes were also

produced
Understanding this result involves exploring
genetic recombination, the production of
offspring with combinations of traits differing
from either parent
Genetic Recombination and

Linkage
The genetic findings of Mendel and Morgan
relate to the chromosomal basis of
recombination
Recombination of Unlinked Genes:

Independent Assortment of Chromosomes
Mendel observed that combinations of traits in
some offspring differ from either parent
Offspring with a phenotype matching one of the
parental phenotypes are called parental types
Offspring with nonparental phenotypes (new
combinations of traits) are called recombinant
types, or recombinants
A 50% frequency of recombination is observed
for any two genes on different chromosomes
Fig. 15-UN2
Gametes from yellow-round

heterozygous parent (YyRr)
Gametes from greenwrinkled homozygous

recessive parent ( yyrr)
YR
yr
Yr
yR
YyRr
yyrr
Yyrr
yyRr
yr
Parentaltype
offspring
Recombinant
offspring
Recombination of Linked Genes: Crossing

Over
Morgan discovered that genes can be linked,
but the linkage was incomplete, as evident
from recombinant phenotypes
Morgan proposed that some process must
sometimes break the physical connection
between genes on the same chromosome
That mechanism was the crossing over of
homologous chromosomes
Animation: Crossing Over
Fig. 15-10
Testcross
parents
Gray body, normal wings

(F1 dihybrid)
Replication
of chromosomes
Meiosis I
Black body, vestigial wings

(double mutant)
b+vg+
bvg
bvg
bvg
b+vg+
bvg
b+vg+
bvg
bvg
bvg
bvg
bvg
b+vg+
Meiosis I and II
b+vg
bvg+
bvg
Meiosis II
Recombinant
chromosomes
Eggs
Testcross
offspring
b+vg+
965
Wild type
(gray-normal)
bvg
944
Blackvestigial
b+vg
206
Grayvestigial
bvg+
185
Blacknormal
bvg
b+vg+
bvg
b vg
b vg
bvg
bvg
bvg
bvg
Parental-type offspring Recombinant offspring

391 recombinants
Recombination
100 = 17%
=
frequency
2,300 total offspring
Sperm
Replication
of chromosomes
Fig. 15-10a
Testcross
parents
Black body, vestigial wings

(double mutant)
Gray body, normal wings

(F1 dihybrid)
Replication
of chromosomes
Meiosis I
b+vg+
bvg
bvg
bvg
b+vg+
bvg
b+vg+
bvg
bvg
bvg
bvg
bvg
b+vg+
Meiosis I and II
b+vg
bvg+
bvg
Meiosis II
Recombinant
chromosomes
b+vg+
b+vg
bvg
Eggs
bvg+
bvg
Sperm
Replication
of chromosomes
Fig. 15-10b
Recombinant
chromosomes
Eggs
Testcross
offspring
b+vg+
bvg
b+vg
bvg+
944
Wild type
Black(gray-normal) vestigial
206
Grayvestigial
185
Blacknormal
965
b+vg+
bvg
b+vg
bvg+
bvg
bvg
bvg
bvg
Parental-type offspring
Recombinant offspring
391 recombinants
Recombination
100 = 17%
=
frequency
2,300 total offspring
bvg
Sperm
Mapping the Distance Between Genes Using

Recombination Data: Scientific Inquiry
Alfred Sturtevant, one of Morgans students,
constructed a genetic map, an ordered list of
the genetic loci along a particular chromosome
Sturtevant predicted that the farther apart two
genes are, the higher the probability that a
crossover will occur between them and
therefore the higher the recombination
frequency
A linkage map is a genetic map of a

chromosome based on recombination
frequencies
Distances between genes can be expressed as
map units; one map unit, or centimorgan,
represents a 1% recombination frequency
Map units indicate relative distance and order,
not precise locations of genes
Fig. 15-11
RESULTS
Recombination
frequencies
9%
Chromosome
9.5%
17%
cn
vg
Genes that are far apart on the same

chromosome can have a recombination
frequency near 50%
Such genes are physically linked, but
genetically unlinked, and behave as if found on
different chromosomes
Sturtevant used recombination frequencies to

make linkage maps of fruit fly genes
Using methods like chromosomal banding,
geneticists can develop cytogenetic maps of
chromosomes
Cytogenetic maps indicate the positions of
genes with respect to chromosomal features
Fig. 15-12
Short
aristae
Long aristae
(appendages
on head)
Mutant phenotypes
Black
body
48.5
Gray
body
Cinnabar Vestigial
eyes
wings
57.5
Red
eyes
67.0
Normal
wings
Wild-type phenotypes
Brown
eyes
104.5
Red
eyes
Concept 15.4: Alterations of chromosome

number or structure cause some genetic disorders
Large-scale chromosomal alterations often
lead to spontaneous abortions (miscarriages)
or cause a variety of developmental disorders
Abnormal Chromosome
Number
In nondisjunction, pairs of homologous
chromosomes do not separate normally during
meiosis
As a result, one gamete receives two of the
same type of chromosome, and another
gamete receives no copy
Fig. 15-13-1
Meiosis I
Nondisjunction
(a) Nondisjunction of homologous

chromosomes in meiosis I
(b) Nondisjunction of sister

chromatids in meiosis II
Fig. 15-13-2
Meiosis I
Nondisjunction
Meiosis II
Nondisjunction


Fig. 15-13-3
Meiosis I
Nondisjunction
Meiosis II
Nondisjunction
Gametes
n+1
n+1
n1
n1
n+1
n1
Number of chromosomes


Aneuploidy results from the fertilization of

gametes in which nondisjunction occurred
Offspring with this condition have an abnormal
number of a particular chromosome
A monosomic zygote has only one copy of a

particular chromosome
A trisomic zygote has three copies of a
particular chromosome
Polyploidy is a condition in which an organism

has more than two complete sets of
chromosomes
Triploidy (3n) is three sets of chromosomes
Tetraploidy (4n) is four sets of chromosomes
Polyploidy is common in plants, but not

animals
Polyploids are more normal in appearance
than aneuploids
Fig. 15-14
Alterations of Chromosome
Structure
Breakage of a chromosome can lead to four
types of changes in chromosome structure:
Deletion removes a chromosomal segment
Duplication repeats a segment
Inversion reverses a segment within a
chromosome
Translocation moves a segment from one
chromosome to another
Fig. 15-15
(a)
(b)
(c)
(d)
A B C D E
F G H
A B C D E
F G H
A B C D E
F G H
A B C D E
F G H
Deletion
Duplication
A B C E
F G H
A B C B C D E
Inversion
A D C B E
F G H
M N O C D E
Reciprocal
translocation
M N O P Q
F G H
A B P Q
F G H
Human Disorders Due to Chromosomal

Alterations
Alterations of chromosome number and
structure are associated with some serious
disorders
Some types of aneuploidy appear to upset the
genetic balance less than others, resulting in
individuals surviving to birth and beyond
These surviving individuals have a set of
symptoms, or syndrome, characteristic of the
type of aneuploidy
Down Syndrome (Trisomy

21)
Down syndrome is an aneuploid condition
that results from three copies of chromosome
21
It affects about one out of every 700 children
born in the United States
The frequency of Down syndrome increases
with the age of the mother, a correlation that
has not been explained
Fig. 15-16
Fig. 15-16a
Fig. 15-16b
Aneuploidy of Sex
Chromosomes
Nondisjunction of sex chromosomes produces
a variety of aneuploid conditions
Klinefelter syndrome is the result of an extra
chromosome in a male, producing XXY
individuals
Monosomy X, called Turner syndrome,
produces X0 females, who are sterile; it is the
only known viable monosomy in humans
Disorders Caused by Structurally Altered

Chromosomes
The syndrome cri du chat (cry of the cat),
results from a specific deletion in chromosome
5
A child born with this syndrome is mentally
retarded and has a catlike cry; individuals
usually die in infancy or early childhood
Certain cancers, including chronic
myelogenous leukemia (CML), are caused by
translocations of chromosomes
Fig. 15-17
Normal chromosome 9
Normal chromosome 22
Reciprocal
translocation
Translocated chromosome 9
Translocated chromosome 22
(Philadelphia chromosome)
Concept 15.5: Some inheritance patterns are

exceptions to the standard chromosome theory
There are two normal exceptions to Mendelian
genetics
One exception involves genes located in the
nucleus, and the other exception involves
genes located outside the nucleus
Genomic
Imprinting
For a few mammalian traits, the phenotype
depends on which parent passed along the
alleles for those traits
Such variation in phenotype is called genomic
imprinting
Genomic imprinting involves the silencing of
certain genes that are stamped with an
imprint during gamete production
Fig. 15-18
Paternal
chromosome
Normal Igf2 allele

is expressed
Maternal
chromosome
Normal Igf2 allele
is not expressed
Wild-type mouse
(normal size)
(a) Homozygote
Mutant Igf2 allele
inherited from mother
Normal size mouse

(wild type)
Mutant Igf2 allele

inherited from father
Dwarf mouse
(mutant)
Normal Igf2 allele

is expressed
Mutant Igf2 allele

is expressed
Mutant Igf2 allele

is not expressed
Normal Igf2 allele

is not expressed
(b) Heterozygotes
Fig. 15-18a
Paternal
chromosome
Normal Igf2 allele

is expressed
Maternal
chromosome
Normal Igf2 allele
is not expressed
(a) Homozygote
Wild-type mouse
(normal size)
Fig. 15-18b
Mutant Igf2 allele

inherited from mother
Normal size mouse

(wild type)
Mutant Igf2 allele

inherited from father
Dwarf mouse
(mutant)
Normal Igf2 allele

is expressed
Mutant Igf2 allele

is expressed
Mutant Igf2 allele

is not expressed
Normal Igf2 allele

is not expressed
(b) Heterozygotes
It appears that imprinting is the result of the

methylation (addition of CH3) of DNA
Genomic imprinting is thought to affect only a
small fraction of mammalian genes
Most imprinted genes are critical for embryonic
development
Fig. 15-UN3
Inheritance of Organelle
Genes
Extranuclear genes (or cytoplasmic genes) are
genes found in organelles in the cytoplasm
Mitochondria, chloroplasts, and other plant
plastids carry small circular DNA molecules
Extranuclear genes are inherited maternally
because the zygotes cytoplasm comes from
the egg
The first evidence of extranuclear genes came
from studies on the inheritance of yellow or
white patches on leaves of an otherwise green
plant
Fig. 15-19
Some defects in mitochondrial genes prevent

cells from making enough ATP and result in
diseases that affect the muscular and nervous
systems
For example, mitochondrial myopathy and
Lebers hereditary optic neuropathy
Fig. 15-UN4
P generation D
gametes
Sperm
C
B
A
Egg
c
b
a
f
The alleles of unlinked
genes are either on
separate chromosomes
(such as d and e) or so
far apart on the same
chromosome (c and f )
that they assort
independently.
This F1 cell has 2n = 6

chromosomes and is
heterozygous for all six
genes shown (AaBbCcDdEeFf ).
Red = maternal; blue = paternal.
D
Each chromosome
has hundreds or
thousands of genes.
Four (A,B,C,F) are
shown on this one.
C
B
A
F
d
E
cb
Genes on the same chromosome whose alleles are so

close together that they do
not assort independently
(such as a,b, and c) are said
to be linked.
Fig. 15-UN5
Fig. 15-UN6
Fig. 15-UN7
Fig. 15-UN8
Fig. 15-UN9
You should now be able

to:
1. Explain the chromosomal theory of
inheritance and its discovery
2. Explain why sex-linked diseases are more
common in human males than females
3. Distinguish between sex-linked genes and
linked genes
4. Explain how meiosis accounts for
recombinant phenotypes
5. Explain how linkage maps are constructed
6. Explain how nondisjunction can lead to

aneuploidy
7. Define trisomy, triploidy, and polyploidy
8. Distinguish among deletions, duplications,
inversions, and translocations
9. Explain genomic imprinting
10.Explain why extranuclear genes are not
inherited in a Mendelian fashion

Unit 4 Chapter 15

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Chapter

Overview: Locating Genes Along

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Concept 15.1: Mendelian inheritance has its

The behavior of chromosomes during meiosis

All F1 plants produce

All F1 plants produce

All F1 plants produce

Morgans Experimental Evidence:

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Morgans Choice of Experimental

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Morgan noted wild type, or normal,

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Correlating Behavior of a Genes Alleles with

Morgan determined that the white-eyed mutant

Concept 15.2: Sex-linked genes exhibit

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

The Chromosomal Basis of

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Females are XX, and males are XY

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

(b) The X-0 system

(c) The Z-W system

(d) The haplo-diploid system

(a) The X-Y system

(b) The X-0 system

(c) The Z-W system

(d) The haplo-diploid system

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Sex-linked genes follow specific patterns of

Sex-linked recessive disorders are much more

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Eggs XN XNXN XNY

Some disorders caused by recessive alleles on

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Cell division and

Concept 15.3: Linked genes tend to be inherited

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

How Linkage Affects

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

b+bvg+vg bbvgvg b+bvgvg bbvg+vg

b+bvg+vg bbvgvg b+bvgvg bbvg+vg

If genes are located on the same chromosome and

Morgan found that body color and wing size

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

However, nonparental phenotypes were also

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Genetic Recombination and

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Recombination of Unlinked Genes:

Gametes from yellow-round

Gametes from greenwrinkled homozygous

Recombination of Linked Genes: Crossing

Gray body, normal wings

Black body, vestigial wings

Parental-type offspring Recombinant offspring

Black body, vestigial wings

Gray body, normal wings

Mapping the Distance Between Genes Using

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

A linkage map is a genetic map of a

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings