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ChapterFour

4
EukaryoticChromosome
MappingbyRecombination
BASICPROBLEMS
1.

Youperformthefollowingcrossandaretoldthatthetwogenesare10m.u.
apart.
AB/abab/ab
Amongtheirprogeny,10percentshouldberecombinant(Ab/abandaB/ab)
and90percentshouldbeparental(AB/ab and ab/ab).Therefore, AB/ab
shouldrepresent1/2oftheparentalsor45percent.

2.

P
F1
F2

Ad/AdaD/aD
Ad/aD
1Ad/Ad
phenotype:Ad
2Ad/aD phenotype:AD
1aD/aD phenotype:aD

3.

RS/rsRS/rs

gametes

1/ (10.35)
2
1/ (10.35)
2
1/ (0.35)
2
1/ (0.35)
2

F1genotypes

0.1056RS/RS

RS
rs
Rs
rS
0.1138rs/rS

55

56ChapterFour
0.1056rs/rs
0.2113RS/rs
0.1138RS/rS
0.1138RS/Rs

0.1138rs/Rs
0.0306Rs/Rs
0.0306rS/rS
0.0613Rs/rS

F1phenotypes 0.6058RS
0.1056rs
0.1444Rs
0.1444rS
4.

Thecrossis E/eF/f e/ef/f.Ifindependentassortmentexists,theprogeny


shouldbeina1:1:1:1ratio,whichisnotobserved.Therefore,thereislinkage.
EfandeFarerecombinantsequalingonethirdoftheprogeny.Thetwogenes
are33.3mapunits(m.u.)apart.
RF=100%1/3=33.3%

5.

Becauseonlyparentaltypesarerecovered,thetwogenesmustbetightlylinked
andrecombinationmustbeveryrare.Knowinghowmanyprogenywerelooked
atwouldgiveanindicationofhowclosethegenesare.

6.

TheproblemstatesthatafemalethatisA/aB/bistestcrossed.Ifthegenesare
unlinked,theyshouldassortindependentlyandthefourprogenyclassesshould
bepresentinroughlyequalproportions.Thisisclearlynotthecase.TheA/a
B/banda/ab/bclasses(theparentals)aremuchmorecommonthantheA/a
b/b and a/aB/b classes(therecombinants).Thetwogenesareonthesame
chromosomeandare10mapunitsapart.
RF=100%(46+54)/1000=10%
A

b
10m.u.

7.

ThecrossisA/Ab/ba/aB/B.TheF1wouldbeA/aB/b.
a. If the genes are unlinked, all four progeny classes from the test cross
(includinga/a;b/b)wouldequal25percent.
b. Withcompletelylinkedgenes,theF1 wouldproduceonly Ab and aB
gametes. Thus, there would be a 0 percent chance of having a b/a b
progenyfromatestcrossofthisF1.

ChapterFour
c. Ifthetwogenesarelinkedand10mapunitsapart,10percentofthetest
crossprogenyshouldberecombinants.SincetheF1isAb/aB,abisoneof
therecombinantclasses(ABbeingtheother)andshouldequal 1/2ofthe
totalrecombinantsor5percent.
8.

d. 12percent(seepartc)
Meiosis is occurring in an organism that is C d/c D, ultimately producing
haploidspores.TheparentalgenotypesareCdandcD,inequalfrequency.The
recombinanttypesareCDandcd,inequalfrequency.Eightmapunitsmeans8
percentrecombinants.Thus,CDandcdwilleachbepresentatafrequencyof
4percent,andCdandcDwilleachbepresentatafrequencyof(100%8%)/2
=46%.
a. 4percent
b. 4percent
c. 46percent
d. 8percent

9.

Toanswerthisquestion,youmustrealizethat
(1)Onechiasmainvolvestwoofthefourchromatidsofthehomologouspairso
if 16 percent of the meioses have one chiasma, it will lead to 8 percent
recombinantsobservedintheprogeny(onehalfofthechromosomesofsucha
meiosisarestillparental),and
(2)HalfoftherecombinantswillbeBr,sothecorrectansweris4percent.

10. UnpackingtheProblem
1. Thereisnocorrectdrawing;anywilldo.Pollenfromthetasselsisplaced
onthesilksofthefemales.TheseedsaretheF1cornkernels.
2. The+salllookthesamebecausetheysignifywildtypeforeachgene.The
informationisgiveninaspecificorder,whichpreventsconfusion,atleast
initially.However,asyouworktheproblem,whichmayrequireyouto
reorderthegenes,errorscancreepintoyourworkifyoudonotmakesure
thatyoureorderthegenesforeachgenotypeinexactlythesameway.You
mayfinditeasiertowritethecompletegenotype,p+insteadof+,toavoid
confusion.
3. Thephenotypeispurpleleavesandbrownmidrifftoseeds.Inotherwords,
thetwocolorsrefertodifferentpartsoftheorganism.
4. Thereisnosignificanceintheoriginalsequenceofthedata.

57

58ChapterFour
5. Atesterisahomozygousrecessiveforallgenesbeingstudied.Itisusedso
thatthemeioticproductsintheorganismbeingtestedcanbeseendirectly
inthephenotypeoftheprogeny.
6. Theprogenyphenotypesallowyoutoinferthegenotypesoftheplants.For
example, gre standsforgreen,thephenotypeof p+/; sen standsfor
virussensitive,thephenotypeof v+/;and pla standsforplainseed,
the phenotype of b+/.Inthis test cross, all progeny have at least one
recessiveallelesothegresenplaprogenyareactuallyp+/pv+/vb+/b.
7. Gametes refers to the gametes of the two purebreeding parents. F1
gametesreferstothegametesproducedbythecompletelyheterozygousF 1
progeny.Theyindicatewhethercrossingoverandindependentassortment
haveoccurred.Inthiscase,becausethereiseitherindependentassortment
orcrossingover,orboth,thedataindicatethatthethreegenesarenotso
tightlylinkedthatzerorecombinationoccurred.
8. ThemainfocusismeiosisoccurringintheF1parent.
9. Thegametesfromthetesterarenotshownbecausetheycontributenothing
tothephenotypicdifferencesseenintheprogeny.
10. Eightphenotypicclassesareexpectedforthreeautosomalgenes,whether
ornottheyarelinked,whenallthreegeneshavesimpledominantrecessive
relationshipsamongtheiralleles.Thegeneralformulaforthenumberof
expectedphenotypesis2n,wherenisthenumberofgenesbeingstudied.
11. If the three genes were on separate chromosomes, the expectation is a
1:1:1:1:1:1:1:1ratio.
12. Thefourclassesofdatacorrespondtotheparentals(largest),twogroupsof
singlecrossovers(intermediate),anddoublecrossovers(smallest).
13. Bycomparingtheparentalswiththedoublecrossovers,geneordercanbe
determined.Thegeneinthemiddleflipswithrespecttothetwoflanking
genesinthedoublecrossoverprogeny.Inthiscase,oneparentalis +++
andonedoublecrossoverisp++.Thisindicatesthatthegeneforleafcolor
(p)isinthemiddle.
14. Ifonlytwoofthethreegenesarelinked,thedatacanstillbegrouped,but
thegroupingwilldifferfromthatmentionedin(12)above.Inthissituation,
theunlinkedgenewillshowindependentassortmentwiththetwolinked
genes. There will be one class composed of four phenotypes in
approximatelyequalfrequency,whichcombinedwilltotalmorethanhalf
the progeny. A second class will be composed of four phenotypes in
approximatelyequalfrequency,andthecombinedtotalwillbelessthan
halftheprogeny.Forexample,ifthecrosswereab/++;c/+ab/ab;

ChapterFour
c/c, then the parental class (more frequent class) would have four
components:abc,ab+,++c,and+++.Therecombinantclasswould
bea+c,a++,+bc,and+b+.
15. Pointreferstolocus.Theusagedoesnotimplylinkage,butratheratesting
forpossiblelinkage.Afourpointtestcrosswouldlooklikethefollowing:
a/+b/+c/+d/+a/ab/bc/cd/d.
16. A recombinant referstoanindividualwhohasallelesinheritedfromtwo
differentgrandparents,bothofwhomweretheparentsoftheindividuals
heterozygousparent.Anotherwaytothinkaboutthistermisthatinthe
recombinant individuals heterozygous parent, recombination took place
amongthegenesthatwereinheritedfromhisorherparents.Inthiscase,
therecombinationtookplaceintheF1andtherecombinantsareamongthe
F2progeny.
17. Therecombinantforcolumnsrefertospecificgenepairsandprogeny
thatexhibitrecombinationbetweenthosegenepairs.
18. There are three recombinant for columns because three genes can be
groupedinthreedifferentgenepairs.
19. R refers to recombinant progeny, and they are determined by reference
backtotheparentsoftheirheterozygousparent.
20. Column totals indicate the number of progeny that experience
crossingoverbetweenthespecificgenepairs.Theyareusedtocalculate
mapunitsbetweenthetwogenes.
21. Thediagnostictestforlinkageisarecombinationfrequencyoflessthan50
percent.
22. A map unit represents 1 percent crossingover and is the same as a
centimorgan.
23. Inthetester,recombinationcannotbedetectedinthegametecontribution
totheprogenybecausethetesterishomozygous.TheF1individualshave
genotypes fixed by their parents homozygous state and, again,
recombination cannotbedetected inthem, simplybecausetheir parents
werehomozygous.
24. Interference I = 1 coefficient of coincidence = 1 (observed double
crossovers/expected double crossovers). The expected double crossovers
areequalto p(frequencyofcrossingoverinthefirstregion,inthiscase
between v and p) p(frequency of crossingover in the second region,
betweenpandb)numberofprogeny.Theprobabilityofcrossingoveris
equaltomapunitsconvertedbacktopercentage.

59

60ChapterFour
25. Ifthethreegenesarenotalllinked,theninterferencecannotbecalculated.
26. Agreatdealofworkisrequiredtoobtain10,000progenyincornbecause
eachseedonacobrepresentsoneprogeny.Eachcobmaycontainasmany
as200seeds.Whileseedcharacteristicscanbeassessedatthecobstage,
forothercharacteristics,eachseedmustbeplantedseparatelyandassessed
aftergerminationandgrowth.Thebookkeepingtaskisalsoenormous.
SolutiontotheProblem
a. Thethreegenesarelinked.
b. Comparingtheparentals(mostfrequent)withthedoublecrossovers(least
frequent),thegeneorderisvpb.Therewere2200recombinantsbetweenv
andp,and1500betweenpandb.Thegeneralformulaformapunitsis
m.u.=100%(numberofrecombinants)/totalnumberofprogeny
Therefore,themapunitsbetweenvandp=100%(2200)/10,000=22m.u.,
andthemapunitsbetweenpandb=100%(1500)/10,000=15m.u.
Themapis
v
p
b
22m.u.

15m.u.

c. I=1observeddoublecrossovers/expecteddoublecrossovers
=1132/(0.22)(0.15)(10,000)
=10.4=0.6
11. P

abc/abca+b+c+/a+b+c+

F1

a+b+c+/abca+b+c+/abc

F2

1364
365
87
84
47
44
5
4

a+b+c+
abc
abc+
a+b+c
ab+c+
a+bc
ab+c
a+bc+

Thisproblemissomewhatsimplifiedbythefactthatrecombinationdoesnot
occur in male Drosophila. Also, only progeny that received the a b c
chromosomefromthemalewillbedistinguishableamongtheF2progeny.

ChapterFour
a. Becauseyoucannotdistinguishbetween a+ b+ c+/a+ b+ c+ and a+ b+
c+/abc,usethefrequencyofabc/abctoestimatethefrequencyofa+
b+c+(parental)gametesfromthefemale.
parentals
COab:
CObc:
DCO:

730
91
171
9
1001

(2365)
(ab+c+,a+bc=47+44)
(abc+,a+b+c=87+84)
(a+bc+,ab+c=4+5)

ab:100%(91+9)/1001=10m.u.
bc:100%(171+9)/1001=18m.u.
b. Coefficientofcoincidence=(observedDCO)/(expectedDCO)
=9/[(0.1)(0.18)(1001)]=0.5
12.a. Bycomparingthetwomostfrequentclasses(parentals:anbr++,an+br
)totheleastfrequentclasses(DCO:an+br+,anbr+),thegeneorder
canbedetermined.Thegeneinthemiddleswitcheswithrespecttothe
othertwo(theorderisanbr).Nowthecrossescanbewrittenfully.
P

an+br+/an+br+an+br/an+br

F1

an+br/an+br+anbr/anbr

F2

355
339
88
55
21
17
2
2
879

b.

an:
br:

an+br+/anbr
an+br/anbr
an++br+/anbr
anbr/anbr
an+br+/anbr
an+br/anbr
an++br/anbr
anbr+/anbr

parental
parental
COan
COan
CObr
CObr
DCO
DCO

100%(88+55+2+2)/879=16.72m.u.
100%(21+17+2+2)/879=4.78m.u.
an

16.72m.u.

br
4.78m.u.

c. Interference=1(observedDCO/expectedDCO)
=14/(0.1672)(0.0478)(879)=0.431

61

62ChapterFour
13. Bycomparingthemostfrequentclasses(parental:+vlg,b++)withtheleast
frequentclasses(DCO:+++,bvlg)thegeneordercanbedetermined.The
geneinthemiddleswitcheswithrespecttotheothertwo,yieldingthefollowing
sequence:vblg.Nowthecrosscanbewritten
P vb+lg/v+blg+vblg/vblg
F1

vb+lg/vblg
v+blg+/vblg
v+blg/vblg
vb+lg+/vblg
v+b+lg/vblg
vblg+/vblg
v+b+lg+/vblg
vblg/vblg

305
275
128
112
74
66
22
18

parental
parental
COblg
COblg
COvb
COvb
DCO
DCO

vb:
blg:

100%(74+66+22+18)/1000=18.0m.u.
100%(128+112+22+18)/1000=28.0m.u.

c.c.

=observedDCO/expectedDCO=(22+18)/(0.28)(0.18)(1000)=0.79

14. Let F =fat, L =longtail, and Fl =flagella. Thegenesequence is F L Fl


(comparemostfrequentwithleastfrequent).Thecrossis
P

FLFl/flflflfl/flfl

F1

398
370
72
67
44
35
9
5

LFl:
FL:

FLFl/flfl
flfl/flfl
FLfl/flfl
flFl/flfl
fLFL/flfl
Flfl/flfl
fLfl/flfl
FlFl/flfl

parental
parental
COLFl
COLFl
COFL
COFL
DCO
DCO

100%(72+67+9+5)/1000=15.3m.u.
100%(44+35+9+5)/1000=9.3m.u.
F

L
9.3m.u.

Fl
15.3m.u.

15.a. Thehypothesisisthatthegenesarenotlinked.Therefore,a1:1:1:1ratiois
expected.

ChapterFour
b. 2 =(5450)2/50+(4750)2/50+(5250)2/50+(4750)2/50
=0.32+0.18+0.08+0.18=0.76
c. With3degreesoffreedom,thepvalueisbetween0.50and0.90.
d. Between50percentand90percentofthetimevaluesthisextremefromthe
predictionwouldbeobtainedbychancealone.
e. Accepttheinitialhypothesis.
f.

Because the 2 value was insignificant, the two genes are assorting
independently.Thegenotypesofallindividualsare
dp+/dp+;e/edp/dp;e+/e+

F1 dp+/dp;e+/e
tester

dp/dp;e/e

progeny

long,ebony
long,gray
short,gray
short,ebony

dp+/dp;e/e
dp+/dp;e+/e
dp/dp;e+/e
dp/dp;e/e

16.a.
Rh+E (R/E/e)
Rhe (r/re/e)

1
Rh+e
(R/re/e)

3
Rh+E
(R/rE/e)

Rhe (r/re/e)
Rh+E (R/rE/e)

5
Rhe
(r/re/e)

1
Rh+E
(R/rE/e)

Rh+e
(R/e/e)

Rh+E
(R/E/e)
b. Yes
c. Dominant
d. As drawn, the pedigree hints at linkage. If unlinked, expect that the
phenotypesofthe10childrenshouldbeina1:1:1:1ratioofRh +E,Rh+e,

63

64ChapterFour
RhE,andRhe.ThereareactuallyfiveRhe,fourRh+E,andoneRh+e.
If linked, this last phenotype would represent a recombinant, and the
distancebetweenthetwogeneswouldbe100%(1/10)=10m.u.However,
thereisjustnotenoughdatatostronglysupportthatconclusion.
17. Assumethereisnolinkage.(Thisisyourhypothesis.Ifitcanberejected,the
genes are linked.) The expected values would be that genotypes occur with
equalfrequency.Therearefourgenotypesineachcase(n =4)sothereare3
degreesoffreedom.
2=(observedexpected)2/expected
Cross1: 2=[(310300)2+(315300)2+(287300)2+(288300)2]/300
=2.1266;p>0.50,nonsignificant;hypothesiscannotberejected
Cross2: 2=[(3630)2+(3830)2+(2330)2+(2330)2]/300
=6.6;p>0.10,nonsignificant;hypothesiscannotberejected
Cross3: 2=[(360300)2+(380300)2+(230300)2+(230300)2]/300
=66.0;p<0.005,significant;hypothesismustberejected
Cross4: 2=[(7460)2+(7260)2+(5060)2+(4460)2]/300
=11.60;p<0.01,significant;hypothesismustberejected
18.a. Bothdisordersmustberecessivetoyieldthepatternsofinheritancethatare
observed. Notice that only males are affected, strongly suggesting X
linkage for both disorders. In the first pedigree there is a 100 percent
correlationbetweenthepresenceorabsenceofbothdisorders,indicating
closelinkage.Inthesecondpedigree,thepresenceandabsenceofboth
disorders are inversely correlated, again indicating linkage. In the first
pedigree, the two defective alleles must be cis within the heterozygous
femalestoshow100percentlinkageintheaffectedmales,whileinthe
second pedigree the two defective alleles must be trans within the
heterozygousfemales.
b. andc. Let a standfortheallelegivingrisetosteroidsulfatasedeficiency
(vertical bar) and b stand for the allele giving rise to ornithine
transcarbamylase deficiency (horizontal bar). Crossing over cannot be
detectedwithoutattachinggenotypestothepedigrees.Whenthisisdone,it
canbeseenthatcrossingoverneednotoccurineitherofthepedigreesto
giverisetotheobservations.

ChapterFour

Firstpedigree:

ab/Y

AB/

AB/ab

AB/Y

AB/ab

AB/Y

ab/Y

AB/Y

ab/Y

AB/

ab/Y

AB/Y

Secondpedigree:

aB/Y

AB/Y

aB/Y

Ab/Ab

Ab/Y

Ab/B

Ab/Y

Ab/Y

19.a.

Notethatonlymalesareaffectedbybothdisorders.Thissuggeststhatboth
areXlinkedrecessivedisorders.UsingpforprotanandPfornonprotan,
anddfordeutanandDfornondeutan,theinferredgenotypesarelistedon
thepedigreebelow.TheYchromosomeisshown,buttheXisrepresented
bytheallelescarried.

b.

IndividualII2musthaveinheritedbothdisordersinthetransconfiguration
(onseparatechromosomes).Therefore,individualIII2inheritedbothtraits
as the result of recombination (crossingover) between his mothers X
chromosomes.

65

66ChapterFour
I

II

pD/Y

PD/Y

Pd/pD

Pd/Y

Pd/Y

PD/Y

PD/

pd/Y

PD/

Pd/Y

pD/Y

III
c.

Pd/PD

Because both genes are Xlinked, this represents crossingover. The


progenysizeistoosmalltogiveareliableestimateofrecombination.

20.a.andb.Again,thebestwaytodeterminewhetherthereislinkageisthrough
chisquareanalysis,whichindicatesthatitishighlyunlikelythatthethree
genes assort independently. To determine linkage by simple inspection,
look at gene pairs. Because this is a testcross, independent assortment
predictsa1:1:1:1ratio.
Comparingshrunkenandwhite,thefrequenciesare
++
(113+4)/total
swh
(116+2)/total
+wh
(2708+626)/total
s+
(2538+601)/total
Thereisnotindependentassortmentbetweenshrunkenandwhite,which
meansthatthereislinkage.
Comparingshrunkenandwaxy,thefrequenciesare
++
swa
+wa
s+

(626+4)/total
(601+2)/total
(2708+113)/total
(2538+1160/total

Thereisnotindependentassortmentbetweenshrunkenandwaxy,which
meansthatthereislinkage.
Comparingwhiteandwaxy,thefrequenciesare
++
whwa
wh+
+wa

(2538+4)/total
(2708+2)/total
(626+116)/total
(601+113)/total

There is not independent assortment between waxy and white, which


meansthatthereislinkage.

ChapterFour

Becauseallthreegenesarelinked,thestrainsmustbe+s+/wh+waand
wh s wa/wh s wa (compare most frequent, parentals, to least frequent,
doublecrossovers,toobtainthegeneorder).Thecrosscanbewrittenas
P

+s+/wh+wawhswa/whswa

F1 asinproblem
Crossoversbetweenwhiteandshrunkenandshrunkenandwaxyare
113
116
4
2
235

601
626
4
2
1233

Dividingbythetotalnumberofprogenyandmultiplyingby100percent
yieldsthefollowingmap:
white

shrunken
3.5m.u.

c.

Interference

waxy
18.4m.u.

=1(observeddoublecrossovers/expecteddouble
crossovers)
=16/(0.035)(0.184)(6,708)=0.86

67

68ChapterFour
21.a. Theresultsofthiscrossindicateindependentassortmentofthetwogenes.
Thismightbediagrammedas

A a a
B

Bb

50%

50%
MEIOSISI

OR
A

a a
B

a a
b

and

and
MEIOSISII

A
B

a
b

A
b

b. Theresultsofthiscrossindicatethatthetwogenesarelinkedand10m.u.
apart.Further,therecessiveallelesareinrepulsioninthedihybrid(Cd/cD
cd/cd).Thismightbediagrammedas

ChapterFour

C Cc
d

C Cc

dD D

dD D

80%

20%
MEIOSISI

OR
C

D D

and

and
MEIOSISII

recombinants

22.a. Ifthegenesareunlinked,thecrossbecomes
P

hyg/hyg;her/herhyg+/hyg+;her+/her+

hyg+/hyg;her+/herhyg+/hyg;her+/her

F2

9/
16
3/
16
3/
16
1/
16

hyg+/;her+/
hyg+/;her/her
hyg/hyg;her+/
hyg/hyg;her/her

Soonly1/16(or6.25percent)oftheseedswouldbeexpectedtogerminate.

69

70ChapterFour
b.andc. No.Morethantwicetheexpectedseedsgerminatedsoassumethe
genesarelinked.Thecrossthenbecomes
P

hygher/hygherhyg+her+/hyg+her+

hyg+her+/hygherhyg+her+/hygher

F2 13percent

hygher/hygher

Because this class represents the combination of two parental


chromosomes,itisequalto
p(hygher)p(hygher)=(1/2parentals)2=0.13
and
parentals=0.72
sorecombinants=10.72=0.28
Therefore,atestcrossofhyg+her+/hyghershouldgive
36%
36%
14%
14%

hyg+her+/hygher
hygher/hygher
hyg+her/hygher
hygher+/hygher

and36percentoftheprogenyshouldgrow(thehygher/hygherclass).
23.

Comparingindependentassortment(RF=0.5)toarecombinationfrequencyof
34percent(RF=0.34)
RF

0.5

0.34
P
0.25
0.33
P
0.25
0.33
R
0.25
0.17
R
0.25
0.17
Theprobability ofobtaining theseresults,assumingindependent assortment,
willbeequalto
0.250.250.250.250.250.25B=0.00024B(whereB=number
ofpossiblebirthordersforfourparentalandtworecombinantprogeny)
ForanRF=0.34,theprobabilityofthisresultwillbeequalto
0.330.170.330.330.170.33B=0.00034B
Theratioofthetwo=0.00034B/0.00024B=1.42andtheLod=0.15

ChapterFour
24.a. Meiosis I (crossingover has occurred between all genes and their
centromeres)
b. Impossible
c. MeiosisI(crossingoverhasoccurredbetweengeneBanditscentromere)
d. MeiosisI
e. MeiosisII
f.

MeiosisII(crossingoverhasoccurredbetweengenesAandBandtheir
centromeres)

g. MeiosisII
h. Impossible
i.

Mitosis

j.

Impossible

25.a.

al2+
al2+
al2
al2
al2+
al2+
al2
al2

al2
al2
al2+
al2+
al2
al2
al2+
al2+

b. The8percentvaluecanbeusedtocalculatethedistancebetweenthegene
andthecentromere.Thatdistanceis1/2thepercentageofseconddivision
segregation,or4percent.
26.a. arg6al2,arg6al2+,arg6+al2+,arg6+al2+
b.

arg6+al2+,arg6+al2,arg6al2+,arg6al2

c. arg6+al2,arg6+al2,arg6al2+,arg6al2+
27.

Theformulaforthisproblemis(i)=emmi/i!wherem=2andi=0,1,or2.
a. (0)=e220/0!=e2=0.135or13.5%

71

72ChapterFour
b. (1)=e221/1!=e2(2)=0.27or27%
c. (2)=e222/2!=e2(2)=0.27or27%
28. UnpackingtheProblem
1. Fungiaregenerallyhaploid.
2. Therearefourpairs,oreightascospores,ineachascus.Onememberof
eachpairispresentedinthedata.
3. Amatingtypeinfungiisanalogoustosexinhumans,inthatthemating
typesoftwoorganismsmustdifferinordertohaveamatingthatproduces
progeny. Mating type is determined experimentally simply by seeing if
progenyresultfromspecificcrosses.
4. Thematingtypes A and a donotindicatedominanceandrecessiveness.
Theysimplysymbolizethematingtypedifference.
5. arg1indicatesthattheorganismrequiresarginineforgrowth.Testingfor
the genotype involves isolating nutritional mutants and then seeing if
argininesupplementationwillallowforgrowth.
6. arg1+ indicates that the organism is wild type and does not require
supplementalarginineforgrowth.
7. Wild type refers to the common form of an organism in its natural
population.
8. Mutantmeansthat,forthetraitbeingstudied,anorganismdiffersfromthe
wildtype.
9. Theactualfunctionoftheallelesinthisproblemdoesnotmatterinsolving
theproblem.
10. Lineartetradanalysisreferstothefactthattheascosporesineachascusare
inalineararrangementthatreflects theorderinwhichthetwomeiotic
divisionsoccurredtoproducethem.Bytrackingtraitsandcorrelatingthem
withposition,itispossibletodetectcrossingovereventsthatoccurredat
thetetrad(fourstrand,homologouspairing)stagepriortothetwomeiotic
divisions.
11. Lineartetradanalysisallowsforthemappingofcentromeresinrelationto
genes,whichcannotbedonewithunorderedtetradanalysis.
12. AcrossismadeinNeurosporabyplacingthetwoorganismsinthesame
testtubeorPetridishandallowingthemtogrow.Gametesdevelopand

ChapterFour
fertilization, followed by meiosis, mitosis, and ascus formation, occurs.
Theasciareisolated,andtheascosporesaredissectedoutofthemwiththe
aidofamicroscope.Theascushasanoctad,oreightspores,withinit,and
thesporesarearrangedinfour(tetrad)pairs.
13. MeiosisoccursimmediatelyfollowingfertilizationinNeurospora.
typeA
(1n)

gamete

gamete

typea
(1n)

fertilization(2n)

meiosis

mitosis

ascospores(1n)
14. Meiosisproducedtheascosporesthatwereanalyzed.
15. ThecrossisAarg1aarg1+.
16. Although there are eight ascospores, they occur in pairs. Each pair
represents one chromatid of the originally paired chromosomes. By
convention,bothmembersofapairarerepresentedbyasinglegenotype.
17. The seven classes represent the seven types of outcomes. The specific
outcomescanbeclassifiedasfollows:
Class

7
outcome
PD
NPD T
T
PD
NPD T
A/a
I
I
I
II
II
II
II
+
arg1 /arg1
I
I
II
I
II
II
II
wherePD=parentalditype,NPD=nonparentalditype,T=tetratype,I=
firstdivisionsegregation,andII=seconddivisionsegregation.
Other classes can be detected, but they indicate the same underlying
process.Forexample,thefollowingthreeasciareequivalent.

73

74ChapterFour
1

3

+
Aarg
Aarg
Aarg+
Aarg+
Aarg
Aarg
aarg
aarg
aarg+
aarg+
aarg+
aarg
Inthefirstascus,acrossoveroccurredbetweenchromatids2and3,while
inthesecondascusitoccurredbetweenchromatids1and3,andinthethird
ascusthecrossoverwasbetweenchromatids1and4.Afourthequivalent
ascus would contain a crossover between chromatids 2 and 3. All four
indicateacrossoverbetweenthesecondgeneanditscentromereandallare
tetratypes.
18. Thisisexemplifiedintheanswerto(17)above.
19. Theclassisidenticalwithclass1intheproblem,butinverted.
20. Linkagearrangementreferstotherelativepositionsofthetwogenesand
thecentromerealongthelengthofthechromosome.
21. A genetic interval refers to the region between two loci, whose size is
measuredinmapunits.
22. Itisnotknownwhetherthetwolociareonseparatechromosomesorareon
thesamechromosome.Thegeneralformulaforcalculatingthedistanceof
alocustoitscentromereistomeasurethepercentageoftetradsthatshow
seconddivisionsegregationpatternsforthatlocusanddividebytwo.
23. Recall that there are eight ascospores per ascus. By inspection, the
frequencyofrecombinantAarg1+ascosporesis4(125)+2(100)+2(36)
+4(4)+2(6)=800.Thereisalsothereciprocalrecombinantgenotype a
arg1.
24. Class1isparental;class2isnonparentalditype.Becausetheyoccurat
equalfrequencies,thetwogenesarenotlinked.
SolutiontotheProblem
a.

ThecrossisAarg1aarg1+.Usetheclassificationofasciinpart(17)
above. First decide if the two genes are linked by using the formula
PD>>NPD,whenthegenesarelinked,whilePD=NPDwhentheyarenot
linked.PD=127+2=129andNPD=125+4=129,whichmeansthat
thetwogenesarenotlinked.Alternatively,
RF

=100%(1/2T+NPD)/totalasci

ChapterFour
=100%[(1/2)(100+36+6)+(125+4)]/400=50%.
Nextcalculatethedistancebetweeneachgeneanditscentromereusingthe
formulaRF=100%(1/2numberoftetradsexhibitingMIIsegregation)/(total
numberofasci).
Acentromere =100%(1/2)(36+2+4+6)/400
=100%(24/400)=6m.u.
arg+centromere=100%(1/2)(100+2+4+6)/400
=100%(56/400)=14m.u.
6m.u.

14m.u.

arg

b. Class6canbeobtainedifasinglecrossoveroccurredbetweenchromatids
2and3betweeneachgeneanditscentromere.

29. Beforebeginningthisproblem,classifyallasciasPD,NPD,orTanddetermine
whetherthereisMIorMIIsegregationforeachgene:

type
genea
geneb

Ascitype

PD
NPD T
T
PD
NPD T
I
I
I
II
II
II
II
I
I
II
I
II
II
II

75

76ChapterFour
If PD >> NPD, linkage is indicated. The distance between a gene and its
centromere=100%(1/2)(MII)/total.Thedistancebetweentwogenes=100%
(1/2T+NPD)/total.
Cross1:PD=NPDandRF=50%;thegenesarenotlinked.
acentromere:100%(1/2)(0)/100=0m.u.Geneaisclosetothecentromere.
bcentromere:100%(1/2)(32)/100=16m.u.
Cross2:PD>>NPD;thegenesarelinked.
ab:100%[1/2(15)+1]/100=8.5m.u.
acentromere:100%(1/2)(0)/100=0m.u.Geneaisclosetothecentromere.
bcentromere:100%(1/2)(15)/100=7.5m.u.
Cross3:PD>>NPD;thegenesarelinked.
ab:100%[1/2(40)+3]/100=23m.u.
acentromere:100%(1/2)(2)/100=1m.u.
bcentromere:100%(1/2)(40+2)/100=21m.u.
Cross4:PD>>NPD;thegenesarelinked.
ab:100%[1/2(20)+1]/100=11m.u.
acentromere:100%(1/2)(10)/100=5m.u.
bcentromere:100%(1/2)(18+8+1)/100=13.5m.u.
Cross5:PD=NPD(andRF=49%);thegenesarenotlinked.
acentromere:100%(1/2)(22+8+10+20)/99=30.3m.u.
bcentromere:100%(1/2)(24+8+10+20)/99=31.3m.u.
Thesevaluesareapproachingthe67percenttheoreticallimitoflociexhibiting
MIIpatternsofsegregationandshouldbeconsideredcautiously.
Cross6:PD>>NPD;thegenesarelinked.
ab:100%[1/2(1+3+4)+0]/100=4m.u.
acentromere:100%(1/2)(3+61+4)/100=34m.u.
bcentromere:100%(1/2)(1+61+4)/100=33m.u.
These values are at the 67 percent theoretical limit of loci exhibiting MII
patternsofsegregationandthereforebothlocicanbeconsideredunlinkedtothe
centromere.
Cross7:PD>>NPD;thegenesarelinked.
ab:100%[1/2(3+2)+0]/100=2.5m.u.
acentromere:100%(1/2)(2)/100=1m.u.
bcentromere:100%(1/2)(3)/100=1.5m.u.

ChapterFour

Cross8:PD=NPD;thegenesarenotlinked.
acentromere:100%(1/2)(22+12+11+22)/100=33.5m.u.
bcentromere:100%(1/2)(20+12+11+22)/100=32.5m.u.
Sameascross5.
Cross9:PD>>NPD;thegenesarelinked.
ab:100%[1/2(10+18+2)+1]/100=16m.u.
acentromere:100%(1/2)(18+1+2)/100.=10.5m.u.
bcentromere:100%(1/2)(10+1+2)/100=6.5m.u.
Cross10:PD=NPD;thegenesarenotlinked.
acentromere:100%(1/2)(60+1+2+5)/100=34m.u.
bcentromere:100%(1/2)(2+1+2+5)/100=5m.u.
a
>50m.u.

or
or

b
5m.u.

b
5m.u.

a
>50m.u.

a
5m.u.

>50m.u.

Cross11:PD=NPD;thegenesarenotlinked.
acentromere:100%(1/2)(0)/100=0m.u.
bcentromere:100%(1/2)(0)/100=0m.u.
30. Thenumberofrecombinants isequaltoNPD+ 1/2T.Theuncorrectedmap
distanceisbasedonRF=(NPD+ 1/2T)/total.Thecorrectedmapdistance=
50(T+6NPD)/total.
Cross1:
recombinantfrequency =4%+1/2(45%)=26.5%
uncorrectedmapdistance=[4%+1/2(45%)]/100%=26.5m.u.
correctedmapdistance =50[45%+6(4%)]/100%=34.5m.u.
Cross2:
recombinantfrequency =2%+1/2(34%)=19%
uncorrectedmapdistance=[2%+1/2(34%)]/100%=19m.u.
correctedmapdistance =50[34%+6(2%)]/100%=29m.u.

77

78ChapterFour
Cross3:
recombinantfrequency =5%+1/2(50%)=30%
uncorrectedmapdistance=[5%+1/2(50%)]/100%=30m.u.
correctedmapdistance =50[50%+6(5%)]/100%=40m.u.

CHALLENGINGPROBLEMS
31.a. Allofthesegenesarelinked.Todeterminethis,eachgenepairisexamined
separately.Forexample,areAandBlinked?
AB=140+305=445
ab=145+310=455
aB=42+6=48
Ab=43+9=52
Conclusion:thetwogenesarelinkedand10m.u.apart.
AreAandDlinked?
AD=0
ad=0
Ad=43+140+9+305=497
aD=42+145+6+310=503
Conclusion:thetwogenesshownorecombinationandatthisresolution,
are0m.u.apart.
AreBandClinked?
BC=42+140=182
bc=43+145=188
Bc=6+305=311
bC=9+310=319
Conclusion:thetwogenesarelinkedand37m.u.apart.
AreCandDlinked?
CD=42+310=350
cd=43+305=348
Cd=140+9=149
cD=145+6=151
Conclusion:thetwogenesarelinkedand30m.u.apart.Therefore,allfour
genesarelinked.

ChapterFour
b.andc. Because A and D shownorecombination,firstrewritetheprogeny
omittingDandd(oromittingAanda).
aBC
Abc
ABC
abc
aBc
AbC
ABc
abC

42
43
140
145
6
9
305
310
1000

Notethattheprogenynowlooklikethoseofatypicalthreepointtestcross,
withABcandabCtheparentaltypes(mostfrequent)andaBcandAbC
thedoublerecombinants(leastfrequent).ThegeneorderisBAC.Thisis
determined either by the map distances or by comparing double
recombinantswiththeparentals;thegenethatswitchesinreferencewith
theothertwoisthegeneinthecenter(BAcBac,baCbAC).
Next,rewritetheprogenyagain,thistimeputtingthegenesintheproper
order,andclassifytheprogeny.
BaC
bAc
BAC
bac
Bac
bAC
BAc
baC

42
43
140
145
6
9
305
310

COAB
COAB
COAC
COAC
DCO
DCO
parental
parental

Toconstructthemapofthesegenes,usethefollowingformula:
distancebetweentwogenes=(100%)(numberofsingleCO+numberofDCO)
totalnumberofprogeny
FortheAtoBdistance
=(100%)(42+43+6+9)=10m.u.
1000
FortheAtoCdistance
=(100%)(140+145+6+9)=30m.u.
1000
Themapis

79

80ChapterFour

A,D
10m.u.

C
30m.u.

Theparentalchromosomesactuallywere B(A,d)c/b(a,D)C,wherethe
parenthesesindicatethattheorderofthegeneswithinisunknown.
d. Interference=1(observedDCO/expectedDCO)
=1(6+9)/[(0.10)(0.30)(1000)]
=115/30=0.5
32. Theverbaldescriptionindicatesthefollowingcrossandresult:
P N/A/n/nO/O
F1

N/nA/ON/nA/O

Theresultsindicatelinkage,sothecrossandresultscanberewritten:
P NA/nO/nO
F1

NA/nONA/nO

F2

66%
16%
9%
9%

NA/orN/A
nO/nO
nA/n
NO/O

Onlyonegenotypeisfullyknown:16percent nO/nO,acombinationoftwo
parentalgametes.Thefrequencyoftwoparentalgametescomingtogetheristhe
frequency of the first times the frequency of the second. Therefore, the
frequencyofeach nO gameteisthesquarerootof0.16,or0.4.Withinan
organismthetwoparental gametes occurinequal frequency.Therefore, the
frequency of N A is also 0.4. The parental total is 0.8, leaving 0.2 for all
recombinants.Therefore,NOandnAoccuratafrequencyof0.1each.Thetwo
genesare20m.u.apart.
33. a.andb.Thedataindicatethattheprogenymaleshaveadifferentphenotype
thanthefemales.Therefore,allthegenesareontheXchromosome.The
twomostfrequentphenotypesinthemalesindicatethegenotypesoftheX
chromosomesinthefemale,andthetwoleastfrequentphenotypesinthe
males indicate the gene order. Data from only the males are used to
determinemapdistances.Thecrossis:

ChapterFour
P

c.

xzy+/x+z+yx+z+y+/Y

F1males
430
xzy+/Y
parental
441
x+z+y/Y
parental
39
xzy/Y COzy
30
x+z+y+/Y
COzy
+
+
32
x zy /Y
COxz
+
27
xz y/Y
COxz
1
x+zy/Y
DCO
+
+
0
xz y /Y
DCO
zy:100%(39+30+1)/1000=7.0m.u.
xz:100%(32+27+1)/1000=6.0m.u.
c.c.

=observedDCO/expectedDCO
=1/[(0.06)(0.07)(1000)]=0.238

34. Thedatagivenforeachofthethreepointtestcrossescanbeusedtodetermine
thegeneorderbyrealizingthattherarestrecombinantclassesaretheresultof
doublecrossoverevents.Bycomparingthesechromosomestotheparental
types,theallelesthathaveswitchedrepresentthegeneinthemiddle.
Forexample,in(1),themostcommonphenotypes(+++andabc)represent
theparentalallelecombinations.Comparingthesetotherarestphenotypesof
thisdataset(+bcanda++)indicatesthattheageneisrecombinantandmust
beinthemiddle.Thegeneorderisbac.
For(2),+bcanda++(theparentals)shouldbecomparedto+++andabc
(therarestrecombinants)toindicatethattheageneisinthemiddle.Thegene
orderisbac.
For(3),compare+b+anda+cwithab+and++c,whichgivesthegene
orderbac.
For(4),compare++candab+with+++andabc,whichgivesthegene
orderacb.
For(5),compare+++andabcwith++candab+,whichgivesthegene
orderacb.
35. Thegeneorderisacbd.
Recombinationbetweenaandcoccurredatafrequencyof
100%(139+3+121+2)/(669+139+3+121+2+2,280+653+2,215)

81

82ChapterFour
=100%(265/6,082)=4.36%
Recombinationbetweenbandcincross1occurredatafrequencyof
100%(669+3+2+653)/(669+139+3+121+2+2,280+653+2,215)
=100%(1,327/6,082)=21.82%
Recombinationbetweenbandcincross2occurredatafrequencyof
100%(8+14+153+141)/(8+441+90+376+14+153+64+141)
=100%(316/1,287)=24.55%
Thedifferencebetweenthetwocalculated distances between b and c isnot
surprising because each set of data would not be expected to yield exactly
identical results. Also, many more offspring were analyzed in cross 1.
Combined,thedistancewouldbe
100%[(316+1,327)/(1,287+6,082)]=22.3%
Recombinationbetweenbanddoccurredatafrequencyof
100%(8+90+14+64)/(8+441+90+376+14+153+64+141)
=100%(176/1,287)=13.68%
Thegeneralmapis:
a

4.4m.u.

22.3m.u.

d
13.7m.u.

36. Part(a)ofthisproblemissolvedtwoways,onceinthestandardway,onceina
waythatemphasizesamoremathematicalapproach.
Thecrossis
P

PAR/PARpar/par

F1

PAR/parpar/par,athreepointtestcross

a. InordertofindwhatproportionwillhavetheVulcanphenotypeforall
three characteristics, we must determine the frequency of parentals.
Crossingoveroccurs15percentofthetimebetweenPandA,whichmeans
itdoesnotoccur85percentofthetime.Crossingoveroccurs20percentof
thetimebetweenAandR,whichmeansthatitdoesnotoccur80percentof
thetime.
p(nocrossoverbetweeneithergene)
=p(nocrossoverbetweenPandA)p(nocrossoverbetweenAandR)
=(0.85)(0.80)=0.68

ChapterFour
HalftheparentalsareVulcan,sotheproportionthatarecompletelyVulcan
is1/2(0.68)=0.34
Mathematicalmethod
Numberofparentals=1(singleCOindividualsDCOindividuals)
=1{[0.15+0.202(0.15)(0.20)](0.15)(0.20)}=0.68
Because half the parentals are Earth alleles and half are Vulcan, the
frequencyofchildrenwithallthreeVulcancharacteristicsis 1/2(0.68)=
0.34
b. Sameasabove,0.34
c. ToyieldVulcanearsandheartsandEarthadrenals,acrossovermustoccur
in both regions, producing double crossovers. The frequency of Vulcan
ears and hearts and Earth adrenals will be half the DCOs, or 1/2(0.15)
(0.20)=0.015
d. ToyieldVulcanearsandanEarthheartandadrenals,asinglecrossover
mustoccurbetweenPandA,andnocrossovercanoccurbetweenAandR.
Thefrequencywillbe
p(COPA)p(noCOAR)=(0.15)(0.80)=0.12
Ofthese, 1/2 are Par and1/2 are pAR.Therefore,theproportionwith
VulcancarsandanEarthheartandadrenalsis0.06
37.a.

Toobtainaplantthatis abc/abc fromselfingof Abc/aBC,both


gametesmustbederivedfromacrossoverbetweenAandB.Thefrequency
oftheabcgameteis
1/ p(COAB)p(noCOBC)=1/ (0.20)(0.70)=0.07
2
2

Therefore,thefrequencyofthehomozygousplantwillbe(0.07)2=0.0049
b. ThecrossisAbc/aBCabc/abc.
Tocalculatetheprogenyfrequencies,notethattheparentalsareequaltoall
thosethatdidnotexperienceacrossover.Mathematicallythiscanbestated
as
parentals

=p(noCOAB)p(noCOBC)
=(0.80)(0.70)=0.56

Becauseeachparentalshouldberepresentedequally

83

84ChapterFour

Abc=1/2(0.56)=0.28
aBC=1/2(0.56)=0.28
Ascalculatedabove,thefrequencyoftheabcgameteis
1/ p(COAB)p(noCOBC)=1/ (0.20)(0.70)=0.07
2
2

asisthefrequencyofABC.
ThefrequencyoftheAbCgameteis
1/ p(COBC)p(noCOAB)=1/ (0.30)(0.80)=0.12
2
2

asisthefrequencyofaBc.
Finally,thefrequencyoftheABcgameteis
1/ p(COAB)p(COBC)=1/ (0.20)(0.30)=0.03
2
2

asisthefrequencyofabC.
Sofor1,000progeny,theexpectedresultsare
Abc
aBC
ABC
abc
AbC
aBc
ABc
abC

280
280
70
70
120
120
30
30

c. Interference=1observedDCO/expectedDCO
0.2=1observedDCO/(0.20)(0.30)
observedDCO=(0.20)(0.30)(0.20)(0.20)(0.30)=0.048
TheABdistance=20%=100%[p(COAB)+p(DCO)].
Therefore,p(COAB)=0.200.048=0.152
Similarly,theBCdistance=30%=100%[p(COBC)+p(DCO)]
Therefore,p(COBC)=0.300.048=0.252
Thep(parental)=1p(COAB)p(COBC)p(observedDCO)
=10.1520.2520.048=0.548

ChapterFour

Sofor1,000progeny,theexpectedresultsare
Abc
aBC
ABC
abc
AbC
aBc
ABc
abC

274
274
76
76
126
126
24
24

38.a. Blue sclerotic (B) appears to be an autosomal dominant disorder.


Hemophilia(h)appearstobeanXlinkedrecessivedisorder.
b. IftheindividualsinthepedigreearenumberedasgenerationsIthroughIV
andtheindividuals ineachgeneration arenumberedclockwise,starting
fromthetoprighthandportionofthepedigree,theirgenotypesare

I:b/b;H/h,B/b;H/Y

II:B/b;H/Y,B/b;H/Y,b/b;H/Y,B/b;H/h,b/b;H/Y,B/b;H/h,B/b;
H/h,B/b;H/,b/b;H/
III:b/b;H/,B/b;H/,b/b;h/Y,b/b;H/Y,B/b;H/Y,B/b;H/,B/b;
H/Y,B/b;h/Y,B/b;H/,b/b;H/Y,B/b;H/,b/b;H/Y,B/b;H/,B/b;
H/Y,B/b;h/Y,b/b;H/Y,b/b;H/Y,b/b;H/,b/b;H/Y,b/b;H/Y,B/b;
H/,B/b;H/Y,B/b;h/Y
IV:b/b;H/,B/b;H/,B/b;H/,b/b;H/h,b/b;H/h,b/b;H/Y,b/b;H/H,
b/b;H/Y,b/b;H/h,b/b;H/H,b/b;H/H,b/b;H/Y,b/b;H/Y,b/b;H/H,
b/b;H/Y,b/b;H/Y,B/b;H/Y,b/b;H/Y,b/b;H/,b/b;H/Y,b/b;H/Y,
b/b;H/,b/b;H/H,b/b;H/,b/b;H/,b/b;H/Y,b/b;H/Y,b/b;H/Y,
b/b;H/h,B/b;H/h,B/b;H/Y,b/b;H/Y,B/b;H/Y,b/b;H/h
c. Thereisnoevidenceoflinkagebetweenthesetwodisorders.Becauseof
the modes of inheritance for these two genes, no linkage would be
expected.
d. Thetwogenesexhibitindependentassortment.
e. No individual could be considered intrachromosomally recombinant.
However,anumbershowinterchromosomalrecombination,forexample,
allindividualsingenerationIIIthathavebothdisorders.

85

86ChapterFour
39. Ifh=hemophiliaandb=colorblindness,thegenotypesforindividualsinthe
pedigreecanbewrittenas

HB/hb

Hb/Y

hb/Y

HB/Y

HB/hb

HB/Y

HB/Y

hb/Y

Hb/B Hb/b

The mother of the two women in question would produce the following
gametes:
0.45
0.45
0.05
0.05

HB
hb
hB
Hb

WomanIII4canbeeither Hb/HB (0.45chance)or Hb/hB (0.05chance),


becauseshereceivedBfromhermother.IfsheisHb/hB[0.05/(0.45+0.05)=
0.10chance],shewillproducetheparentalandrecombinantgameteswiththe
sameprobabilitiesashermother.Thus,herchildhasa45percentchanceof
receivinghB,a5percentchanceofreceivinghb,anda50percentchanceof
receiving a Y from his father. The probability that her child will be a
hemophiliacsonis(0.1)(0.5)(0.5)=0.025=2.5percent.
WomanIII5canbeeither Hb/Hb (0.05chance)or Hb/hb (0.45chance),
becauseshereceivedbfromhermother.IfsheisHb/hb[0.45/(0.45+0.05)=
0.90chance],shehasa50percentchanceofpassinghtoherchild,andthereis
a50percentchancethatthechildwillbemale.Theprobabilitythatshewill
haveasonwithhemophiliais(0.9)(0.5)(0.5)=0.225=22.5percent.
40.a. Cross1reducesto
P

A/AB/BD/Da/ab/bd/d

F1

A/aB/bD/da/ab/bd/d

Thetestcrossprogenyindicatethesethreegenesarelinked.
Testcross ABD

316

parental

ChapterFour
progeny

abd
ABd
abD
Abd
aBD
AbD
aBd

314
31
39
130
140
17
13

87

parental
COBD
COBD
COAB
COAB
DCO
DCO

AB: 100%(130+140+17+13)/1000=30m.u.
BD: 100%(31+39+17+13)/1000=10m.u.
Cross2reducesto
P
A/AC/CE/Ea/ac/ce/e
F1

A/aC/cE/ea/ac/ce/e

Thetestcrossprogenyindicatethesethreegenesarelinked.
Testcross ACE
progeny
ace
Ace
aCE
ACe
acE
aCe
AcE

243
237
62
58
155
165
46
34

parental
parental
COAC
COAC
COCE
COCE
DCO
DCO

AC: 100%(62+58+46+34)/1000=20m.u.
CE: 100%(155+165+46+34)/1000=40m.u.
Themapthataccommodatesallthedatais
E

C
40m.u.

A
20m.u.

B
30m.u.

D
10m.u.

b. Interference(I)=1[(observedDCO)/(expectedDCO)]
Forcross1:I=1{30/[(0.30)(0.10)(1000)]}=11=0,nointerference
Forcross2:I=1{80/[(0.20)(0.40)(1000)]}=11=0,nointerference
41.a. ThefirstF1isLH/lhandthesecondislH/Lh.Forprogenythatarelh/lh,
they have received a parental chromosome from the first F1 and a
recombinantchromosomefromthesecondF1.Thegenesare16percent

88ChapterFour
apartsothechanceofaparentalchromosomeis 1/2(10016%)=42%and
thechanceofarecombinantchromosomeis1/2(16%)=8%.
b.

Thechanceofbothevents=42%8%=3.36%
ToobtainLh/lhprogeny,eitheraparentalchromosomefromeachparent
was inherited or a recombinant chromosome from each parent was
inherited.Thetotalprobabilitywillthereforebe
(42%42%)+(8%8%)=(17.6%+0.6%)=18.2%

42.

Crossingoveroccurs8percentofthetimebetween w and s,whichmeansit


doesnotoccur92percentofthetime.Crossingoveroccurs14percentofthe
timebetweensande,whichmeansthatitdoesnotoccur86percentofthetime.
a.andb.Thefrequencyofparentals=p(nocrossoverbetweeneithergene)

or

=p(noCOws)p(noCOse)=(0.92)(0.86)
=0.791
1/ (0.791)=0.396each
2

c.andd.Thefrequencythatwillshowrecombinationbetweenwandsonly
or

=p(COws)p(noCOse)=(0.08)(0.86)=0.069
1/ (0.069)=0.035each
2

e.andf. Thefrequencythatwillshowrecombinationbetweensandeonly
or

=p(COse)p(noCOws)=(0.14)(0.92)=0.128
1/ (0.128)=0.064each
2

g.andh.Thefrequencythatwillshowrecombinationbetween wandsands
ande
or

43.

=p(COws)p(COse)=(0.08)(0.14)=0.011
1/ (0.011)=0.006each
2

Thisproblemisanalogoustomeiosisinorganismsthatformlineartetrads.Let
red=Randblue=r.Thiscannowbecomparedtomeiosisinanorganismthat
isR/r.Thepatterns,theirfrequencies,andthedivisionofsegregationaregiven

ChapterFour
below.Noticethattheprobabilitieschangeaseachball/alleleisselected.This
occurswhenthereissamplingwithoutreplacement.

1/2R

1/2r

1/ 3R 1/1r1/1r=1/6RRrr

firstdivision

1/2R1/1r=1/6RrRr
2/ 3r or
1/2r1/ 1R=1/6RrrR

seconddivision

1/ 3r 1/1R1/1R=1/6rrRR

firstdivision

1/ 2R 1/1r=1/ 6rRRr
2/ 3R or
1/ 2r1/ 1R=1/6RrRr

seconddivision

seconddivision

seconddivision

These results indicate onethird firstdivision segregation and twothirds


seconddivisionsegregation.
44.

Astheproblemsuggests,calculatethefrequenciesofthevariouspossibilities.
Thepercentageoftetradswithoutcrossingoveris88% 80%=70.4%.The
percentageoftetradswithasinglecrossoverinregion(i)andnoneinregion(ii)
is12% 80%=9.6%.Thepercentageoftetradswithasinglecrossoverin
region(ii)andnoneinregion(i)is20% 88%=17.6%andthepercentageof
tetradswithcrossoversinbothregionsis12%20%=2.4%.
Nowworkoutthepatternsofsegregationthatresultineachcase
Fornocrossovers
p

pq
pq
++
++
MIMIPD

Forasinglecrossoverinregion(i)
p

pq
++
pq
++
MIIMIIPD

89

90ChapterFour
Forasinglecrossoverinregion(ii)
p

pq
p+
+q

++
MIMIIT

Fordoublecrossovers,therearefourtypes,allequallylikely
twostrand
p

pq
+q
p+

fourstrand

++
MIIMIT
p+
++
pq

andtwodifferentthreestrand
p

+q
MIIMIT
p+
+q
pq

++
MIIMIIT
pq
++
p+

(a)MIMIPDistheresultofnocrossovers=70.4%

+q
MIIMIIT

ChapterFour

(b)MIMINPDisnotfoundasaresult
(c)MIMIITistheresultofasinglecrossoverinregion(ii)=17.6%
(d)MIIMITistheresultofthetwoandfourstranddoublecrossovers=1.2%
(e)MIIMIIPDistheresultofasinglecrossoverinregion(i)=9.6%
(f)MIIMIINPDisnotfoundasresult
(g)MIIMIITistheresultofboththreestranddoublecrossovers=1.2%
45.a.andb.Thedatasupporttheindependentassortmentoftwogenes(callthem
arg1andarg2).Thecrossbecomesarg1;arg2+arg1+;arg2andthe
resultingtetradsare:
4:0(PD)
arg1;arg2+
arg1;arg2+
arg1+;arg2
arg1+;arg2

3:1(T)
arg1;arg2+
arg1+;arg2
arg1;arg2
arg1+;arg2+

BecausePD=NPD,thegenesareunlinked.

2:2(NPD)
arg1;arg2
arg1;arg2
arg1+;arg2+
arg1+;arg2+

91