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ICS1.
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 Genetics  1.1  
Michael  Deyholos  
30  November,  2010  
 
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Šƒ’–‡”ͳ 


Š› †‘ ‘ˆˆ•’”‹‰ Ž‘‘ Ž‹‡ –Š‡‹” ’ƒ”‡–•ǫ  ˜‡ ƒ…‹‡– ’‡‘’Ž‡ ™‡”‡
ƒ™ƒ”‡–Šƒ––Š‡…Šƒ”ƒ…–‡”‹•–‹…•‘ˆƒ‹†‹˜‹†—ƒŽ’Žƒ–‘”ƒ‹ƒŽ…‘—Ž†„‡
’ƒ••‡† „‡–™‡‡ ‰‡‡”ƒ–‹‘•Ǥ  Š‡› ƒŽ•‘ ‡™ –Šƒ– •‘‡ Š‡”‹–ƒ„Ž‡
…Šƒ”ƒ…–‡”‹•–‹…• ȋ•—…Š ƒ• –Š‡ •‹œ‡ ‘ˆ ˆ”—‹–Ȍ ˜ƒ”‹‡† „‡–™‡‡ ‹†‹˜‹†—ƒŽ•ǡ
ƒ†–Šƒ––Š‡›…‘—Ž†•‡Ž‡…–ˆ‘”–Š‡‘•–ˆƒ˜‘”ƒ„Ž‡–”ƒ‹–•™Š‹Ž‡„”‡‡†‹‰
…”‘’•ƒ†ƒ‹ƒŽ•ǤŠ‡‘…‡’”‡˜ƒŽ‡–ȋ„—–‘™†‹•…”‡†‹–‡†Ȍ…‘…‡’–‘ˆ
blending inheritance ’”‘’‘•‡† –Šƒ– ƒ —†‡ˆ‹‡† ‡••‡…‡ǡ ‹ ‹–•
‡–‹”‡–›ǡ …‘–ƒ‹‡† ƒŽŽ ‘ˆ –Š‡ Š‡”‹–ƒ„Ž‡ ‹ˆ‘”ƒ–‹‘ ˆ‘” ƒ ‹†‹˜‹†—ƒŽǤ
—…ŠŽ‹‡–Š‡‹š‹‰‘ˆ–™‘…‘Ž‘”•‘ˆ’ƒ‹–ǡ‹–™ƒ•–Š‘—‰Š––Šƒ–ƒ–‹‰
…‘„‹‡† –Š‡ ‡••‡…‡• ˆ”‘ ‡ƒ…Š ’ƒ”‡–Ǣ ‘…‡ „Ž‡†‡† –‘‰‡–Š‡”ǡ –Š‡
‹†‹˜‹†—ƒŽ…Šƒ”ƒ…–‡”‹•–‹…•‘ˆ–Š‡’ƒ”‡–•…‘—Ž†‘–„‡•‡’ƒ”ƒ–‡†ƒ‰ƒ‹Ǥ

 

Mendel™ƒ•‘‡‘ˆ–Š‡ˆ‹”•––‘–ƒ‡ƒ•…‹‡–‹ˆ‹…ƒ’’”‘ƒ…Š–‘–Š‡•–—†›‘ˆ
Š‡”‡†‹–›Ǥ  ‡ •–ƒ”–‡† ™‹–Š ™‡ŽŽǦ…Šƒ”ƒ…–‡”‹œ‡† ƒ–‡”‹ƒŽ•ǡ ”‡’‡ƒ–‡† Š‹•
‡š’‡”‹‡–•ƒ›–‹‡•ǡƒ†‡’–…ƒ”‡ˆ—Ž”‡…‘”†•‘ˆŠ‹•‘„•‡”˜ƒ–‹‘•Ǥ
‘” ‡šƒ’Ž‡ǡ ™‘”‹‰ ™‹–Š ’‡ƒ•ǡ ‡†‡Ž •Š‘™‡† –Šƒ– ™Š‹–‡ǦˆŽ‘™‡”‡†
’Žƒ–• …‘—Ž† „‡ ’”‘†—…‡† „› …”‘••‹‰ –™‘ ’—”’Ž‡ǦˆŽ‘™‡”‡† ’Žƒ–•ǡ „—–
‘Ž› ‹ˆ –Š‡ ’—”’Ž‡ǦˆŽ‘™‡”‡† ’Žƒ–• –Š‡•‡Ž˜‡• Šƒ† ƒ– Ž‡ƒ•– ‘‡ ™Š‹–‡Ǧ
ˆŽ‘™‡”‡†’ƒ”‡–ȋ ‹‰ͳǤʹȌǤŠ‹•™ƒ•‡˜‹†‡…‡–Šƒ––Š‡‰‡‡–‹…ˆƒ…–‘”–Šƒ–
’”‘†—…‡†™Š‹–‡ǦˆŽ‘™‡”•Šƒ†not„Ž‡†‡†‹””‡˜‡”•‹„Ž›™‹–Š–Š‡ˆƒ…–‘”ˆ‘”
’—”’Ž‡ǦˆŽ‘™‡”•Ǥ  ‡†‡Žǯ• ‘„•‡”˜ƒ–‹‘• Š‡Ž’‡† –‘ †‹•’”‘˜‡ „Ž‡†‹‰
‹Š‡”‹–ƒ…‡ǡ ‹ ˆƒ˜‘” ‘ˆ ƒ ƒŽ–‡”ƒ–‹˜‡ …‘…‡’– …ƒŽŽ‡† particulate Figure 1.1
inheritanceǡ ‹ ™Š‹…Š Š‡”‡†‹–› ‹• –Š‡ ’”‘†—…– ‘ˆ †‹•…”‡–‡ ˆƒ…–‘”• –Šƒ– Gregor Mendel
…‘–”‘Ž‹†‡’‡†‡––”ƒ‹–•Ǥƒ…ŠŠ‡”‡†‹–ƒ”›ˆƒ…–‘”…‘—Ž†‡š‹•–‹‘‡‘”
‘”‡†‹ˆˆ‡”‡–˜‡”•‹‘•ǡ‘”allelesǤ

‡†‡Žǯ•†‹•…”‡–‡ˆƒ…–‘”•‘ˆŠ‡”‡†‹–›Žƒ–‡”„‡…ƒ‡‘™ƒ•genesǤ
–Š‡‹” ƒ””‘™‡•– †‡ˆ‹‹–‹‘ǡ ‰‡‡• ƒ”‡ ƒ„•–”ƒ…– …‘…‡’–•ǣ —‹–• ‘ˆ


 Š ƒ ’ – ‡ ”  ͳ ȁ1Ǧ2

‹Š‡”‹–ƒ…‡Ǥ  Š‡ …‘‡…–‹‘ „‡–™‡‡ ‰‡‡• ƒ† •—„•–ƒ…‡• Ž‹‡ 

ƒ† …Š”‘‘•‘‡• ™ƒ• ‡•–ƒ„Ž‹•Š‡† Žƒ”‰‡Ž› –Š”‘—‰Š –Š‡ ‡š’‡”‹‡–•
†‡•…”‹„‡†‹–Š‡”‡ƒ‹†‡”‘ˆ–Š‹•…Šƒ’–‡”Ǥ‘™‡˜‡”ǡ‹–‹•™‘”–Š‘–‹‰
–Šƒ– ‡†‡Ž ƒ† ƒ› ”‡•‡ƒ”…Š‡”• ™Š‘ ˆ‘ŽŽ‘™‡† Š‹ ™‡”‡ ƒ„Ž‡ –‘
’”‘˜‹†‡‰”‡ƒ–‹•‹‰Š–•‹–‘„‹‘Ž‘‰›•‹’Ž›„›‘„•‡”˜‹‰–Š‡‹Š‡”‹–ƒ…‡
‘ˆ•’‡…‹ˆ‹…–”ƒ‹–•Ǥ
Figure 1.2 Inheritance of
flower color in peas.
Mendel observed that a
cross between pure
breeding,white and
purple peas (generation
P) produced only
progeny (generation F1)
with purple flowers.
However, white flowered
plant reappeared among
the F2 generation progeny
of a mating between two
F1 plants. The symbols
P, F1 and F2 are
abbreviations for  
parental, first filial, and
second filial generations, 

respectively.
› –Š‡ ‡ƒ”Ž› ͳͻͲͲǯ•ǡ „‹‘…Š‡‹•–• Šƒ† ‹•‘Žƒ–‡† Š—†”‡†• ‘ˆ †‹ˆˆ‡”‡–
…Š‡‹…ƒŽ• ˆ”‘ Ž‹˜‹‰ …‡ŽŽ•Ǥ  Š‹…Š ‘ˆ –Š‡•‡ ™ƒ• –Š‡ ‰‡‡–‹… ƒ–‡”‹ƒŽǫ
”‘–‡‹• •‡‡‡† Ž‹‡ ’”‘‹•‹‰ …ƒ†‹†ƒ–‡•ǡ •‹…‡ –Š‡› ™‡”‡ ƒ„—†ƒ–
ƒ† …‘’Ž‡š ‘Ž‡…—Ž‡•Ǥ ‘™‡˜‡”ǡ ƒ ˆ‡™ ‡› ‡š’‡”‹‡–• Š‡Ž’‡† –‘
’”‘˜‡–Šƒ–ǡ”ƒ–Š‡”–Šƒ’”‘–‡‹ǡ‹•–Š‡‰‡‡–‹…ƒ–‡”‹ƒŽǤ

‹…”‘„‹‘Ž‘‰‹•–• Šƒ† ‹†‡–‹ˆ‹‡† –™‘ •–”ƒ‹• ‘ˆ –Š‡ „ƒ…–‡”‹—

Streptococcus pneumoniaeǤ  ‡ •–”ƒ‹ ȋȌ ’”‘†—…‡† ”‘—‰Š …‘Ž‘‹‡•ǡ
™Š‹Ž‡–Š‡‘–Š‡”ȋȌ™ƒ••‘‘–Šȋ ‹‰ǤͳǤ͵ȌǤ‘”‡‹’‘”–ƒ–Ž›ǡ–Š‡Ǧ–›’‡
„ƒ…–‡”‹ƒ …ƒ—•‡† ˆƒ–ƒŽ ‹ˆ‡…–‹‘•ǡ ™Š‹Ž‡ –Š‡ Ǧ–›’‡ †‹† ‘–Ǥ  Griffith
‘–‹…‡† –Šƒ– •‹’Ž› ‹š‹‰ –Š‡ •–”ƒ‹• –‘‰‡–Š‡” …‘—Ž† –”ƒ•ˆ‘” •‘‡
Ǧ–›’‡ „ƒ…–‡”‹ƒ ‹–‘ •‘‘–Šǡ ’ƒ–Š‘‰‡‹… •–”ƒ‹• ȋ ‹‰Ǥ ͳǤͶȌǤ  Š‹•
–”ƒ•ˆ‘”ƒ–‹‘‘……—””‡†‡˜‡™Š‡–Š‡Ǧ–›’‡•–”ƒ‹•™‡”‡ˆ‹”•–‹ŽŽ‡†
„›Š‡ƒ–ǤŠ—•ǡ•‘‡…‘’‘‡–‘ˆ–Š‡Ǧ–›’‡•–”ƒ‹•…‘–ƒ‹‡†‰‡‡–‹…
‹ˆ‘”ƒ–‹‘–Šƒ–…‘—Ž†„‡–”ƒ•ˆ‡””‡†–‘–Š‡Ǧ–›’‡•–”ƒ‹•Ǥ

Figure 1.3 Colonies of
Rough (top) and Smooth
(bottom) strains of S.
pneumoniae.

Šƒ––›’‡‘ˆ‘Ž‡…—Ž‡ˆ”‘™‹–Š‹–Š‡Ǧ–›’‡…‡ŽŽ•™ƒ•”‡•’‘•‹„Ž‡ˆ‘”
–Š‡ –”ƒ•ˆ‘”ƒ–‹‘ǫ  ‘ ƒ•™‡” –Š‹•ǡ ”‡•‡ƒ”…Š‡”• ƒ‡† Avery,
MacLeod and McCarty •‡’ƒ”ƒ–‡† –Š‡ Ǧ–›’‡ …‡ŽŽ• ‹–‘ ˜ƒ”‹‘—•
…‘’‘‡–•ǡ•—…Šƒ•’”‘–‡‹•ǡ’‘Ž›•ƒ……Šƒ”‹†‡•ǡŽ‹’‹†•ǡƒ†—…Ž‡‹…ƒ…‹†•Ǥ
Ž›–Š‡—…Ž‡‹…ƒ…‹†•ˆ”‘Ǧ–›’‡…‡ŽŽ•™‡”‡ƒ„Ž‡–‘ƒ‡–Š‡Ǧ•–”ƒ‹•
•‘‘–Š ƒ† ˆƒ–ƒŽǤ  —”–Š‡”‘”‡ǡ ™Š‡ …‡ŽŽ—Žƒ” ‡š–”ƒ…–• ‘ˆ Ǧ–›’‡ …‡ŽŽ•
™‡”‡ –”‡ƒ–‡† ™‹–Š ƒ•‡ ȋƒ ‡œ›‡ –Šƒ– †‹‰‡•–• Ȍǡ –Š‡
–”ƒ•ˆ‘”ƒ–‹‘ ƒ„‹Ž‹–› ™ƒ• Ž‘•–Ǥ  Š‡ ”‡•‡ƒ”…Š‡”• –Š‡”‡ˆ‘”‡ …‘…Ž—†‡†
–Šƒ–  ™ƒ• –Š‡ ‰‡‡–‹… ƒ–‡”‹ƒŽǡ ™Š‹…Š ‹ –Š‹• …ƒ•‡ …‘–”‘ŽŽ‡† –Š‡
ƒ’’‡ƒ”ƒ…‡ƒ†’ƒ–Š‘‰‡‹…‹–›‘ˆ–Š‡„ƒ…–‡”‹ƒǤ


 Š ƒ ’ – ‡ ”  ͳ ȁ1Ǧ3


Figure 1.4 Experiments
of Griffith and of Avery ,
MacLeod and McCarty.
R strains of S.
pneumoniae do not cause
lethality. However,
DNA-containing extracts
from pathogenic S strains
are sufficient to make R
strains pathogenic.




—”–Š‡” ‡˜‹†‡…‡ –Šƒ–  ‹• –Š‡ ‰‡‡–‹… ƒ–‡”‹ƒŽ …ƒ‡ ˆ”‘
‡š’‡”‹‡–• …‘†—…–‡† „› Hershey and ChaseǤ  Š‡•‡ ”‡•‡ƒ”…Š‡”•
•–—†‹‡†–Š‡–”ƒ•‹••‹‘‘ˆ‰‡‡–‹…‹ˆ‘”ƒ–‹‘ˆ”‘ƒ˜‹”—•…ƒŽŽ‡†–Š‡
ʹ„ƒ…–‡”‹‘’Šƒ‰‡–‘‹–•Š‘•–„ƒ…–‡”‹—ǡEscherichiacoliȋ ‹‰ǤͳǤͷȌǤ‹‡
ƒŽŽ˜‹”—•‡•ǡʹŠ‹Œƒ…•–Š‡…‡ŽŽ—Žƒ”ƒ…Š‹‡”›‘ˆ‹–•Š‘•––‘ƒ—ˆƒ…–—”‡
‘”‡ ˜‹”—•‡•Ǥ  ‹”—•‡• …‘–ƒ‹ „‘–Š ’”‘–‡‹ ƒ† Ǥ  ‘ †‡–‡”‹‡
™Š‹…Š ‘ˆ –Š‡•‡ –›’‡• ‘ˆ ‘Ž‡…—Ž‡• …‘–ƒ‹‡† –Š‡ ‰‡‡–‹… „Ž—‡’”‹– ˆ‘”
–Š‡ ˜‹”—•ǡ ‡”•Š‡› ƒ† Šƒ•‡ ‰”‡™ ˜‹”ƒŽ …—Ž–—”‡• ‹ –Š‡ ’”‡•‡…‡ ‘ˆ
”ƒ†‹‘ƒ…–‹˜‡‹•‘–‘’‡•‘ˆ‡‹–Š‡”’Š‘•’Š‘”—•ȋ͵ʹȌ‘”•—Ž’Š—”ȋ͵ͷȌǡ™Š‹…Š
‹…‘”’‘”ƒ–‡† –Š‡•‡ ‹•‘–‘’‡• ‹–‘ –Š‡‹”  ƒ† ’”‘–‡‹•ǡ ”‡•’‡…–‹˜‡Ž›
ȋ ‹‰ ͳǤ͸ȌǤ  Š‡ ”‡•‡ƒ”…Š‡”• –Š‡ ‹ˆ‡…–‡† E. coli ™‹–Š –Š‡ ”ƒ†‹‘Žƒ„‡Ž‡† Figure 1.5
Electronmicrograph of
˜‹”—•‡•ǡ ƒ† Ž‘‘‡† –‘ •‡‡ ™Š‡–Š‡” ͵ʹ ‘” ͵ͷ ‡–‡”‡† –Š‡ „ƒ…–‡”‹ƒǤ
T2 bacteriophage on
ˆ–‡” ‡•—”‹‰ –Šƒ– ƒ› ˜‹”—•‡• Šƒ† „‡‡ ”‡‘˜‡† ˆ”‘ –Š‡ •—”ˆƒ…‡ ‘ˆ surface of E. coli
–Š‡ …‡ŽŽ•ǡ –Š‡ ”‡•‡ƒ”…Š‡”• ‘„•‡”˜‡† –Šƒ– ‘Ž› ‹ˆ‡…–‹‘ ™‹–Š ͵ʹ Žƒ„‡Ž‡†
˜‹”—•‡•”‡•—Ž–‡†‹”ƒ†‹‘ƒ…–‹˜‡„ƒ…–‡”‹ƒǤŠ‹•ƒ‰ƒ‹‹†‹…ƒ–‡†–Šƒ–
™ƒ•–Š‡ƒ–‡”‹ƒŽ–Šƒ–…‘–ƒ‹‡†‰‡‡–‹…‹•–”—…–‹‘•Ǥ


 Š ƒ ’ – ‡ ”  ͳ ȁ1Ǧ4

Figure 1.6 When 32P-

labeled phage infects E.
coli, radioactivity is
found only in the bacteria
after the phage are
removed by agitation and
centrifugation. In
contrast, after infection
with 35S-labeled phage,
radioactivity is found
only in the supernatant
that remains after the
bacteria are removed.

 
Š‡‡š’‡”‹‡–•‘ˆ‡”•Š‡› ƒ†Šƒ•‡ǡ˜‡”›ǡ ƒ…‡‘†ǡ ƒ† …ƒ”–›ǡ

”‹ˆˆ‹–Šǡƒ†‘–Š‡”•’”‘˜‡†–Šƒ–™ƒ•–Š‡‰‡‡–‹…ƒ–‡”‹ƒŽǡ„—–˜‡”›
Ž‹––Ž‡™ƒ•‘™ƒ„‘—–‹–••–”—…–—”‡ǤŠ‡WatsonandCrick•‡–‘—–
–‘ †‡–‡”‹‡ –Š‡ •–”—…–—”‡ ‘ˆ  ‹ –Š‡ ͳͻͶͲǯ• –Š‡› ‡™ –Šƒ– 
™ƒ• ƒ†‡ —’ ‘ˆ ˆ‘—” †‹ˆˆ‡”‡– –›’‡• ‘ˆ ‘Ž‡…—Ž‡•ǡ …ƒŽŽ‡† „ƒ•‡•ǡ ‘”
—…Ž‡‘–‹†‡•ǣ ƒ†‡‹‡ ȋȌǡ …›–‘•‹‡ ȋȌǡ –Š›‹‡ ȋȌǡ ‰—ƒ‹‡ ȋ
ȌǤ  Š‡›
ƒŽ•‘ ‡™ ‘ˆ Chargaff’’s Rulesǡ ™Š‹…Š ™ƒ• ƒ •‡– ‘ˆ ‘„•‡”˜ƒ–‹‘• ƒ„‘—–
–Š‡ ”‡Žƒ–‹˜‡ ƒ‘—– ‘ˆ ‡ƒ…Š —…Ž‡‘–‹†‡ –Šƒ– ™ƒ• ’”‡•‡– ‹ ƒŽ‘•– ƒ›
‡š–”ƒ…–‘ˆǤŠƒ”‰ƒˆˆŠƒ†‘„•‡”˜‡†–Šƒ–ˆ‘”ƒ›‰‹˜‡•’‡…‹‡•ǡ–Š‡
ƒ„—†ƒ…‡‘ˆ™ƒ•–Š‡•ƒ‡ƒ•ǡƒ†
™ƒ•–Š‡•ƒ‡ƒ•Ǥ•‹‰‡–ƒŽ
‘†‡Ž• ‘ˆ –Š‡ ‹†‹˜‹†—ƒŽ —…Ž‡‘–‹†‡•ǡ ƒ–•‘ ƒ† ”‹… ™‡”‡ ƒ„Ž‡ –‘
†‡†—…‡ ƒ •–”—…–—”‡ ˆ‘”  –Šƒ– ™ƒ• …‘•‹•–‡– ™‹–Š Šƒ”‰ƒˆˆǯ• —Ž‡•
ƒ† ™‹–Š šǦ”ƒ› …”›•–ƒŽŽ‘‰”ƒ’Š› †ƒ–ƒ –Šƒ– ™ƒ• ‘„–ƒ‹‡† ȋ™‹–Š •‘‡
…‘–”‘˜‡”•›Ȍˆ”‘ƒ‘–Š‡””‡•‡ƒ”…Š‡”ǡ‘•ƒŽ‹† ”ƒŽ‹Ǥ

 ƒ–•‘ ƒ† ”‹…ǯ• ˆƒ‘—• †‘—„Ž‡ Š‡Ž‹šǡ ‡ƒ…Š ‘ˆ –Š‡ –™‘ •–”ƒ†•
…‘–ƒ‹•  „ƒ•‡• …‘‡…–‡† –Š”‘—‰Š …‘˜ƒŽ‡– „‘†• –‘ ƒ •—‰ƒ”Ǧ
’Š‘•’Šƒ–‡ „ƒ…„‘‡ ȋ ‹‰ ͳǤ͹ǡ ͳǤͺȌǤ   ‡…ƒ—•‡ ‘‡ •‹†‡ ‘ˆ ‡ƒ…Š •—‰ƒ”
‘Ž‡…—Ž‡ ‹• ƒŽ™ƒ›• …‘‡…–‡† –‘ –Š‡ ‘’’‘•‹–‡ •‹†‡ ‘ˆ –Š‡ ‡š– •—‰ƒ”
‘Ž‡…—Ž‡ǡ ‡ƒ…Š •–”ƒ† ‘ˆ  Šƒ• ’‘Žƒ”‹–›ǣ –Š‡•‡ ƒ”‡ …ƒŽŽ‡† –Š‡ ͷǯ ȋͷǦ
’”‹‡Ȍ ‡† ƒ† –Š‡ ͵ǯ ȋ͵Ǧ’”‹‡Ȍ ‡†ǡ ‹ ƒ……‘”†ƒ…‡ ™‹–Š –Š‡
‘‡…Žƒ–—”‡ ‘ˆ –Š‡ …ƒ”„‘• ‹ –Š‡ •—‰ƒ”•Ǥ  Š‡ –™‘ •–”ƒ†• ‘ˆ –Š‡
†‘—„Ž‡ Š‡Ž‹š ”— ‹ ƒ–‹Ǧ’ƒ”ƒŽŽ‡Ž ȋ‹Ǥ‡Ǥ ‘’’‘•‹–‡Ȍ †‹”‡…–‹‘•ǡ ™‹–Š –Š‡ ͷǯ
‡†‘ˆ‘‡•–”ƒ†ƒ†Œƒ…‡––‘–Š‡͵ǯ‡†‘ˆ–Š‡‘–Š‡”•–”ƒ†ǤŠ‡†‘—„Ž‡
Figure 1.7 DNA structure Š‡Ž‹šŠƒ•ƒ”‹‰Š–ǦŠƒ†‡†–™‹•–ǡȋ”ƒ–Š‡”–Šƒ–Š‡Ž‡ˆ–ǦŠƒ†‡†–™‹•––Šƒ–‹•
‘ˆ–‡ ”‡’”‡•‡–‡† ‹ ’‘’—Žƒ” ‡†‹ƒȌǤ  Š‡  „ƒ•‡• ‡š–‡† ˆ”‘ –Š‡
„ƒ…„‘‡–‘™ƒ”†•–Š‡…‡–‡”‘ˆ–Š‡Š‡Ž‹šǡ™‹–Šƒ’ƒ‹”‘ˆ„ƒ•‡•ˆ”‘‡ƒ…Š
•–”ƒ† ˆ‘”‹‰ Š›†”‘‰‡ „‘†• –Šƒ– Š‡Ž’ –‘ Š‘Ž† –Š‡ –™‘ •–”ƒ†•



 Š ƒ ’ – ‡ ”  ͳ ȁ1Ǧ5

–‘‰‡–Š‡”Ǥ  †‡” ‘•– …‘†‹–‹‘•ǡ –Š‡ –™‘ •–”ƒ†• ƒ”‡ •Ž‹‰Š–Ž› ‘ˆˆ•‡–ǡ
…”‡ƒ–‹‰ ƒ ƒŒ‘” ‰”‘‘˜‡ ‘ ‘‡ ˆƒ…‡ ‘ˆ –Š‡ †‘—„Ž‡ Š‡Ž‹šǡ ƒ† ƒ ‹‘”
‰”‘‘˜‡‘–Š‡‘–Š‡”Ǥ‡…ƒ—•‡‘ˆ–Š‡•–”—…–—”‡‘ˆ–Š‡„ƒ•‡•ǡ…ƒ‘Ž›
ˆ‘”Š›†”‘‰‡„‘†•™‹–Šǡƒ†
…ƒ‘Ž›ˆ‘”Š›†”‘‰‡„‘†•™‹–Š
 ȋŠ‡…‡ǡ Šƒ”‰ƒˆˆǯ• —Ž‡•ȌǤ  ƒ…Š •–”ƒ† ‹• –Š‡”‡ˆ‘”‡ •ƒ‹† –‘ „‡
…‘’Ž‡‡–ƒ”› –‘ –Š‡ ‘–Š‡”ǡ ƒ† ‡ƒ…Š •–”ƒ† ƒŽ•‘ …‘–ƒ‹• ‡‘—‰Š
‹ˆ‘”ƒ–‹‘ –‘ ”‡’Žƒ…‡ –Š‡ ‘–Š‡”Ǥ   Š‹• ”‡†—†ƒ…› ‹• ‹’‘”–ƒ– ‹
”‡’ƒ‹”‹‰ǡƒ†ƒŽ•‘‹‹–•”‡’Ž‹…ƒ–‹‘Ǥ
Figure 1.8 Chemical
structure of two pairs of
nucleotides in a fragment
of double-stranded DNA.
Sugar, phosphate, and
bases A,C,G,T are
labeled. Hydrogen bonds
between bases on
opposite strands are
shown by dashed lines.
Note that the G-C pair
has more hydrogen bonds
than A-T. The numbering
of carbons within sugars
is indicated by red
numbers. Based on this
numbering the polarity of
each strand is indicated
by the labels 5’’ and 3’’.


  
‘™†‘‡•–Š‡•–”—…–—”‡‘ˆ”‡Žƒ–‡–‘‹Š‡”‹–ƒ…‡‘ˆ„‹‘Ž‘‰‹…ƒŽ–”ƒ‹–•
•—…Š ƒ• –Š‡ ˆŽ‘™‡” …‘Ž‘” ‘ˆ ‡†‡Žǯ• ’‡ƒ•ǫ  Š‡ ƒ•™‡” –‘ –Š‹• Ž‹‡• ‹
™Šƒ–Šƒ•„‡…‘‡‘™ƒ•‘Ž‡…—Žƒ”„‹‘Ž‘‰›ǯ•CentralDogmaǡ™Š‹…Š
•–ƒ–‡• –Šƒ– ‡ƒ…Š ‰‡‡ ‹• ‡…‘†‡† ‹ ǡ ƒ† –Š‡ ƒ• ‡‡†‡†ǡ –Š‹•
‰‡‡–‹… ‹ˆ‘”ƒ–‹‘ ‹• –”ƒ•…”‹„‡† ‹–‘  ƒ† –Š‡ –”ƒ•Žƒ–‡† ‹–‘
’”‘–‡‹Ǥ …‡”–ƒ‹…‹”…—•–ƒ…‡•ǡƒ›ƒŽ•‘„‡…‘˜‡”–‡†–‘
–Š”‘—‰Š ƒ ’”‘…‡•• …ƒŽŽ‡† ”‡˜‡”•‡ –”ƒ•…”‹’–‹‘Ǥ  Š‡ ‘”†‡” ‘ˆ „ƒ•‡• ‹
†‹”‡…–Ž›…‘–”‘Ž•–Š‡‘”†‡”‘ˆƒ‹‘ƒ…‹†•–Šƒ–ƒ‡—’ƒ’”‘–‡‹Ǥ
”‘–‡‹•†‘ ‘•–‘ˆ–Š‡ ™‘”‹ ƒ…‡ŽŽǡƒ†…ƒ–ƒŽ›œ‡–Š‡ˆ‘”ƒ–‹‘ƒ†
„”‡ƒ†‘™ ‘ˆ ƒŽ‘•– ƒŽŽ ‘ˆ –Š‡ ‘Ž‡…—Ž‡• ™‹–Š‹ ƒ ‘”‰ƒ‹•Ǥ   › Figure 1.9 Central
†‹…–ƒ–‹‰–Š‡•–”—…–—”‡‘ˆ‡ƒ…Š’”‘–‡‹ǡƒˆˆ‡…–•–Š‡ˆ—…–‹‘‘ˆ–Šƒ– Dogma of molecular
’”‘–‡‹ǡ ƒ† …ƒ –Š‡”‡„› ƒˆˆ‡…– –Š‡ ‡–‹”‡ ‘”‰ƒ‹•Ǥ   –Š‡ …ƒ•‡ ‘ˆ biology
‡†‡Žǯ• ’‡ƒ•ǡ ’—”’Ž‡ǦˆŽ‘™‡”‡† ’Žƒ–• Šƒ˜‡ ƒ ‰‡‡ –Šƒ– ‡…‘†‡• ƒ
‡œ›‡ –Šƒ– ’”‘†—…‡• ƒ ’—”’Ž‡ ’‹‰‡– ‘Ž‡…—Ž‡Ǥ   –Š‡ ™Š‹–‡Ǧ
ˆŽ‘™‡”‡† ’Žƒ–•ǡ –Š‡  ˆ‘” –Š‹• ‰‡‡ Šƒ• „‡‡ …Šƒ‰‡† •‘ –Šƒ– ‹– ‘
Ž‘‰‡” ‡…‘†‡• ƒ ˆ—…–‹‘ƒŽ ’”‘–‡‹Ǥ  Š‹• ‹• ƒ ‡šƒ’Ž‡ ‘ˆ ƒ ƒ–—”ƒŽ
—–ƒ–‹‘‹ƒ„‹‘…Š‡‹…ƒŽ’ƒ–Š™ƒ›Ǥ


 Š ƒ ’ – ‡ ”  ͳ ȁ1Ǧ6

Š‡…‘’Ž‡–‡•‡–‘ˆ™‹–Š‹–Š‡—…Ž‡—•‘ˆƒ›‘”‰ƒ‹•‹•…ƒŽŽ‡†‹–•
‰‡‘‡Ǥ  ”‰ƒ‡ŽŽ‡• •—…Š ƒ• ‹–‘…Š‘†”‹ƒ ƒ† …ŠŽ‘”‘’Žƒ•–• ƒŽ•‘ Šƒ˜‡
–Š‡‹”‘™‰‡‘‡•ǤŠ‡”‡‹••—”’”‹•‹‰Ž›Ž‹––Ž‡…‘””‡Žƒ–‹‘„‡–™‡‡–Š‡
—…Ž‡ƒ”…‘–‡–‘ˆƒ‰‡‘‡ȋ‹Ǥ‡Ǥ–Š‡cǦvalueȌƒ†–Š‡’Š›•‹…ƒŽ•‹œ‡
‘”…‘’Ž‡š‹–›‘ˆƒ‘”‰ƒ‹•Ǥ ‘”‡šƒ’Ž‡ǡƒ•‹‰Ž‡…‘’›‘ˆ–Š‡Š—ƒ
‰‡‘‡ …‘–ƒ‹• ƒ’’”‘š‹ƒ–‡Ž› ͵ š ͳͲͻ  „ƒ•‡•ǡ ™Š‹Ž‡ ƒ •‹‰Ž‡
™Š‡ƒ– ‰‡‘‡ …‘–ƒ‹• ͳ͹ š ͳͲͻ  „ƒ•‡•Ǥ   Š‹• ƒ’’ƒ”‡– ’ƒ”ƒ†‘š
ȋ…ƒŽŽ‡† –Š‡ …Ǧ˜ƒŽ—‡ ’ƒ”ƒ†‘šȌ …ƒ „‡ ‡š’Žƒ‹‡† „› –Š‡ ˆƒ…– –Šƒ– ‘– ƒŽŽ
 ‡…‘†‡• ‰‡‡•Ǥ   ˆƒ…–ǡ ‹ ƒ› ‘”‰ƒ‹••ǡ ‰‡‡• ƒ”‡ •‡’ƒ”ƒ–‡†
ˆ”‘‡ƒ…Š‘–Š‡”„›Ž‘‰•–”‡–…Š‡•‘ˆ–Šƒ–†‘‘–…‘†‡ˆ‘”ƒ’”‘–‡‹Ǥ
‘‡ ‘ˆ –Š‹• Dz‘Ǧ…‘†‹‰dz  ƒ› „‡ –”ƒ•’‘•‘•ǡ ™Š‹…Š ƒ”‡ ƒ
‹–‡”‡•–‹‰ …Žƒ•• ‘ˆ •‡ŽˆǦ”‡’Ž‹…ƒ–‹‰  ‡Ž‡‡–• †‹•…—••‡† ‹ ‘”‡
†‡–ƒ‹Ž‹ƒ•—„•‡“—‡–…Šƒ’–‡”Ǥ


Table 1.1 Measures of genome size in selected organisms. The DNA content (1C) is shown in millions of
basepairs (Mb). Average gene density is the mean number of non-coding bases (in bp) between genes in the
genome. For eukaryotes, the chromosome number is the chromosomes counted in a gamete (1N) from each
organism.



‡Šƒ˜‡•‡‡ƒŽ”‡ƒ†›–Šƒ–ƒ›‘ˆ–Š‡‰”‡ƒ–ƒ†˜ƒ…‡•‹‰‡‡–‹…•™‡”‡
ƒ†‡ —•‹‰ •’‡…‹‡• –Šƒ– ƒ”‡ ‘– ‡•’‡…‹ƒŽŽ› ‹’‘”–ƒ– ˆ”‘ ƒ ‡†‹…ƒŽǡ
‡…‘‘‹…ǡ ‘” ‡˜‡ ‡…‘Ž‘‰‹…ƒŽ ’‡”•’‡…–‹˜‡Ǥ  
‡‡–‹…‹•–•ǡ ˆ”‘ ‡†‡Ž
‘™ƒ”†•ǡ Šƒ˜‡ —•‡† ‘†‡Ž ‘”‰ƒ‹•• ˆ‘” –Š‡‹” ‡š’‡”‹‡–•Ǥ  ‘†ƒ›ǡ ƒ
•ƒŽŽ—„‡”‘ˆ•’‡…‹‡•ƒ”‡™‹†‡Ž›—•‡†ƒ‘†‡Ž‰‡‡–‹…‘”‰ƒ‹••ǤŽŽ
‘ˆ –Š‡•‡ •’‡…‹‡• Šƒ˜‡ …Šƒ”ƒ…–‡”‹•–‹…• –Šƒ– ƒ‡ –Š‡ ‡ƒ•› –‘ ‰”‘™ ‹
Žƒ”‰‡—„‡”•‹Žƒ„‘”ƒ–‘”‹‡•ǣ–Š‡›ƒ”‡•ƒŽŽǡˆƒ•–‰”‘™‹‰™‹–Šƒ•Š‘”–
‰‡‡”ƒ–‹‘–‹‡ƒ†’”‘†—…‡Ž‘–•‘ˆ’”‘‰‡›ˆ”‘ƒ–‹‰•–Šƒ–…ƒ„‡
‡ƒ•‹Ž› …‘–”‘ŽŽ‡†Ǥ 
‡‡–‹… ‘†‡Ž ‘”‰ƒ‹•• ƒŽ•‘ —•—ƒŽŽ› Šƒ˜‡ •ƒŽŽ
‰‡‘‡•ȋ•ƒŽŽ…Ǧ˜ƒŽ—‡Ȍǡƒ†ƒ”‡†‹’Ž‘‹†ȋ‹Ǥ‡Ǥ…Š”‘‘•‘‡•ƒ”‡’”‡•‡–
‹ ’ƒ‹”•ȌǤ  ‡ƒ•– ȋSaccharomyces cerevisiaeȌ ‹• ƒ ‰‘‘† ‰‡‡”ƒŽ ‘†‡Ž
ˆ‘” –Š‡ „ƒ•‹… ˆ—…–‹‘• ‘ˆ ‡—ƒ”›‘–‹… …‡ŽŽ•Ǥ  Š‡ ”‘—†™‘”ǡ
Caenorhabditis elegans ‹• ƒ —•‡ˆ—Ž ‘†‡Ž ˆ‘” –Š‡ †‡˜‡Ž‘’‡– ‘ˆ



 Š ƒ ’ – ‡ ”  ͳ ȁ1Ǧ7

—Ž–‹…‡ŽŽ—Žƒ”‘”‰ƒ‹••ǡ‹’ƒ”–„‡…ƒ—•‡‹–‹•–”ƒ•’ƒ”‡––Š”‘—‰Š‘—–‹–•
Ž‹ˆ‡…›…Ž‡ǡƒ†‹–•…‡ŽŽ•—†‡”‰‘ƒ™‡ŽŽǦ…Šƒ”ƒ…–‡”‹œ‡†•‡”‹‡•‘ˆ†‹˜‹•‹‘•
–‘ ’”‘†—…‡ –Š‡ ƒ†—Ž– „‘†›Ǥ  Š‡ ˆ”—‹– ˆŽ› ȋDrosophila melanogasterȌ
Šƒ• „‡‡ •–—†‹‡† Ž‘‰‡”ǡ ƒ† ’”‘„ƒ„Ž› ‹ ‘”‡ †‡–ƒ‹Žǡ –Šƒ ƒ› ‘ˆ –Š‡
‘–Š‡” ‰‡‡–‹… ‘†‡Ž ‘”‰ƒ‹•• •–‹ŽŽ ‹ —•‡ǡ ƒ† ‹• ƒ —•‡ˆ—Ž ‘†‡Ž ˆ‘”
•–—†›‹‰†‡˜‡Ž‘’‡–ƒ•™‡ŽŽƒ•’Š›•‹‘Ž‘‰›ƒ†‡˜‡„‡Šƒ˜‹‘—”Ǥ•ƒ
ƒƒŽǡ ‘—•‡ ȋMus musculusȌ ‹• –Š‡ ‘†‡Ž ‘”‰ƒ‹• ‘•– …Ž‘•‡Ž›
”‡Žƒ–‡†–‘Š—ƒ•ǡŠ‘™‡˜‡”•‘‡‘ˆ–Š‡’”ƒ…–‹…ƒŽ†‹ˆˆ‹…—Ž–‹‡•‘ˆ™‘”‹‰
™‹–Š ‹…‡ Ž‡† ”‡•‡ƒ”…Š‡”• ‘”‡ ”‡…‡–Ž› –‘ †‡˜‡Ž‘’ œ‡„”ƒˆ‹•Š ȋDanio
rerioȌ ƒ• ƒ ‰‡‡–‹… ‘†‡Ž ˆ‘” ˜‡”–‡„”ƒ–‡•Ǥ  Ž‹‡ ‹…‡ǡ œ‡„”ƒˆ‹•Š
‡„”›‘• †‡˜‡Ž‘’ ‡š–‡”ƒŽŽ› –‘ –Š‡‹” ‘–Š‡”• ƒ† ƒ”‡ –”ƒ•’ƒ”‡–ǡ
ƒ‹‰ ‹– ‡ƒ•‹‡” –‘ •–—†› –Š‡‹” †‡˜‡Ž‘’‡–Ǥ  ‹ƒŽŽ›ǡ ƒ •ƒŽŽ ™‡‡†ǡ
Arabidopsis thalianaǡ ‹• –Š‡ ‘•– ™‹†‡Ž› •–—†‹‡† ’Žƒ– ‰‡‡–‹… ‘†‡Ž
‘”‰ƒ‹•Ǥ


Š‡ •–—†› ‘ˆ ‰‡‡–‹… ‘†‡Ž ‘”‰ƒ‹•• Šƒ• ‰”‡ƒ–Ž› ‹…”‡ƒ•‡† ‘—” Figure 1.10 Some of the
‘™Ž‡†‰‡ ‘ˆ ‰‡‡–‹…•ǡ ƒ† „‹‘Ž‘‰› ‹ ‰‡‡”ƒŽǤ  ‘†‡Ž ‘”‰ƒ‹•• ƒŽ•‘ most important genetic
Šƒ˜‡‹’‘”–ƒ–‹’Ž‹…ƒ–‹‘•‹‡†‹…ƒŽ”‡•‡ƒ”…ŠǤ ‘”‡šƒ’Ž‡ǡƒ–Ž‡ƒ•– model organisms in use
today. Clockwise from
͹ͷΨ ‘ˆ–Š‡ ƒ’’”‘š‹ƒ–‡Ž›ͳǡͲͲͲ‰‡‡•–Šƒ–Šƒ˜‡„‡‡ ƒ••‘…‹ƒ–‡†™‹–Š
top left: yeast, fruit fly,
•’‡…‹ˆ‹…Š—ƒ†‹•‡ƒ•‡•Šƒ˜‡Š‹‰ŠŽ›•‹‹Žƒ”•‡“—‡…‡•‹„‘–ŠŠ—ƒ• arabidopsis, mouse,
ƒ† D. melanogasterǤ   ˆ‘”ƒ–‹‘ Ž‡ƒ”‡† ˆ”‘ ‘†‡Ž ‘”‰ƒ‹•• roundworm, zebrafish.
ƒ„‘—–’ƒ”–‹…—Žƒ”„‹‘…Š‡‹…ƒŽ’ƒ–Š™ƒ›•…ƒ—•—ƒŽŽ›„‡ƒ’’Ž‹‡†–‘‘–Š‡”
•’‡…‹‡•ǡ•‹…‡–Š‡ƒ‹ˆ‡ƒ–—”‡•‘ˆƒ›„‹‘…Š‡‹…ƒŽ’ƒ–Š™ƒ›•–‡†–‘
„‡•Šƒ”‡†„‡–™‡‡•’‡…‹‡•Ǥ

– ‹• ƒŽ•‘ ’‘••‹„Ž‡ǡ ƒ† •‘‡–‹‡• ‡…‡••ƒ”›ǡ –‘ •–—†› „‹‘Ž‘‰‹…ƒŽ

’”‘…‡••‡•‹‘Ǧ‘†‡Ž‘”‰ƒ‹••Ǥ—ƒ•ǡˆ‘”‡šƒ’Ž‡ǡŠƒ˜‡‘‡‘ˆ
–Š‡…Šƒ”ƒ…–‡”‹•–‹…•‘ˆƒ‘†‡Ž‘”‰ƒ‹•ǡƒ†–Š‡”‡ƒ”‡•‘‡†‹•‡ƒ•‡•‘”
‘–Š‡”–”ƒ‹–•ˆ‘”™Š‹…Š‘…Ž‡ƒ”ƒƒŽ‘‰‡š‹•–•‹‘–Š‡”‘”‰ƒ‹••Ǥ‘‡
‘ˆ–Š‡–‘‘Ž•‘ˆ‰‡‡–‹…ƒƒŽ›•‹•…ƒ„‡ƒ’’Ž‹‡†–‘‘Ǧ‘†‡Ž‘”‰ƒ‹••ǡ
‡•’‡…‹ƒŽŽ› ™‹–Š –Š‡ †‡˜‡Ž‘’‡– ‘ˆ ‡™ –›’‡• ‘ˆ ‰‡‡–‹… ƒ’’‹‰ ƒ†
™Š‘Ž‡‰‡‘‡•‡“—‡…‹‰Ǥ


 Š ƒ ’ – ‡ ”  ͳ ȁ1Ǧ8

̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴


x ‡†‡Ž †‡‘•–”ƒ–‡† –Šƒ– Š‡”‡†‹–› ‹˜‘Ž˜‡† †‹•…”‡–‡ǡ Š‡”‹–ƒ„Ž‡ ˆƒ…–‘”• –Šƒ– ƒˆˆ‡…–‡†
•’‡…‹ˆ‹…–”ƒ‹–•Ǥ

x ‰‡‡…ƒ„‡†‡ˆ‹‡†ƒ„•–”ƒ…–Ž›ƒ•—‹–‘ˆ‹Š‡”‹–ƒ…‡Ǣƒ›‰‡‡–‹…‡š’‡”‹‡–•…ƒ„‡
…‘†—…–‡†™‹–Š‘—–ƒ‘™Ž‡†‰‡‘ˆǤ

x Š‡ƒ„‹Ž‹–›‘ˆˆ”‘„ƒ…–‡”‹ƒƒ†˜‹”—•‡•–‘–”ƒ•ˆ‡”‰‡‡–‹…‹ˆ‘”ƒ–‹‘‹–‘„ƒ…–‡”‹ƒ
Š‡Ž’‡†’”‘˜‡–Šƒ–‹•–Š‡‰‡‡–‹…ƒ–‡”‹ƒŽǤ

x  ‹• ƒ †‘—„Ž‡ Š‡Ž‹š ƒ†‡ ‘ˆ –™‘ ƒ–‹Ǧ’ƒ”ƒŽŽ‡Ž •–”ƒ†• ‘ˆ „ƒ•‡• ‘ ƒ •—‰ƒ”Ǧ’Š‘•’Šƒ–‡
„ƒ…„‘‡Ǥ

x ’‡…‹ˆ‹… „ƒ•‡• ‘ ‘’’‘•‹–‡ •–”ƒ†• ’ƒ‹” –Š”‘—‰Š Š›†”‘‰‡ „‘†‹‰ǡ ‡•—”‹‰
…‘’Ž‡‡–ƒ”‹–›‘ˆ–Š‡•–”ƒ†•Ǥ

x Š‡‡–”ƒŽ‘‰ƒ‡š’Žƒ‹•Š‘™ƒˆˆ‡…–•Š‡”‹–ƒ„Ž‡–”ƒ‹–•Ǥ

x ‘–ƒŽŽ‘ˆ–Š‡‹ƒ‘”‰ƒ‹•…‘–ƒ‹•‰‡‡•Ǥ

x ‘†‡Ž‘”‰ƒ‹••ƒ……‡Ž‡”ƒ–‡–Š‡—•‡‘ˆ‰‡‡–‹…•‹„ƒ•‹…ƒ†ƒ’’Ž‹‡†”‡•‡ƒ”…Š‹„‹‘Ž‘‰›ǡ
ƒ‰”‹…—Ž–—”‡ƒ†‡†‹…‹‡Ǥ


„Ž‡†‹‰‹Š‡”‹–ƒ…‡ „ƒ…–‡”‹‘’Šƒ‰‡ ‡–”ƒŽ‘‰ƒ
’ƒ”–‹…—Žƒ–‡‹Š‡”‹–ƒ…‡ Šƒ”‰ƒˆˆǯ•—Ž‡• –”ƒ•…”‹’–‹‘
‡†‡Ž ƒ–•‘ƒ†”‹… –”ƒ•Žƒ–‹‘
‰‡‡ „ƒ•‡• 
ƒŽŽ‡Ž‡ •—‰ƒ”Ǧ’Š‘•’Šƒ–‡„ƒ…„‘‡ ‰‡‘‡
–”ƒ‹– ƒ–‹Ǧ’ƒ”ƒŽŽ‡Ž …Ǧ˜ƒŽ—‡’ƒ”ƒ†‘š
ǡ ͳǡ ʹ …‘’Ž‡‡–ƒ”› ‘†‡Ž‘”‰ƒ‹•

”‹ˆˆ‹–Š Š›†”‘‰‡„‘† Saccharomycescerevisiae

˜‡”›ǡƒ…‡‘†ǡ…ƒ”–› ‹‘”‰”‘‘˜‡ Caenorhabditiselegans
‡”•Š‡›ƒ†Šƒ•‡ ƒŒ‘”‰”‘‘˜‡ Drosophilamelanogaster
ƒ•‡ ƒ†‡‹‡ Musmusculus
’”‘–‡‹ƒ•‡ …›–‘•‹‡ Daniorerio
͵ͷ –Š›‹‡ Arabidopsisthaliana
͵ʹ ‰—ƒ‹‡ Escherichiacoli


̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴̴



 Š ƒ ’ – ‡ ”  ͳ ȁ1Ǧ9


1.1   ‘™ ™‘—Ž† –Š‡ ”‡•—Ž–• ‘ˆ –Š‡ …”‘•• ‹
‹‰—”‡ ͳǤʹ Šƒ˜‡ „‡‡ †‹ˆˆ‡”‡– ‹ˆ Š‡”‡†‹–›
™‘”‡†–Š”‘—‰Š„Ž‡†‹‰‹Š‡”‹–ƒ…‡”ƒ–Š‡”
–Šƒ’ƒ”–‹…—Žƒ–‡‹Š‡”‹–ƒ…‡ǫ

ƒ††‹–‹‘ƒŽ –‘ –Š‘•‡ •Š‘™ ‹ ‹‰—”‡ ͳǤͶ –‘
†‡‘•–”ƒ–‡ –Šƒ–  ‹• –Š‡ ‰‡‡–‹…
ƒ–‡”‹ƒŽǫ

1.6 ‹•– –Š‡ ‹ˆ‘”ƒ–‹‘ –Šƒ– ƒ–•‘ ƒ†
1.2  ƒ‰‹‡ –Šƒ– ƒ•–”‘ƒ—–• ’”‘˜‹†‡ ›‘—
™‹–Š Ž‹˜‹‰ •ƒ’Ž‡• ‘ˆ —Ž–‹…‡ŽŽ—Žƒ”
‘”‰ƒ‹•• †‹•…‘˜‡”‡† ‘ ƒ‘–Š‡” ’Žƒ‡–Ǥ
Š‡•‡ ‘”‰ƒ‹•• ”‡’”‘†—…‡ ™‹–Š ƒ •Š‘”–
‰‡‡”ƒ–‹‘ –‹‡ǡ „—– ‘–Š‹‰ ‡Ž•‡ ‹• ‘™
ƒ„‘—––Š‡‹”‰‡‡–‹…•Ǥ
a) ‘™ …‘—Ž† ›‘— †‡ˆ‹‡ Žƒ™• ‘ˆ
Š‡”‡†‹–›ˆ‘”–Š‡•‡‘”‰ƒ‹••ǫ
b) ‘™ …‘—Ž† ›‘— †‡–‡”‹‡ ™Šƒ–
‘Ž‡…—Ž‡• ™‹–Š‹ –Š‡•‡ ‘”‰ƒ‹••
…‘–ƒ‹‡†‰‡‡–‹…‹ˆ‘”ƒ–‹‘ǫ
c) ‘—Ž† –Š‡ ‡…Šƒ‹•• ‘ˆ ‰‡‡–‹…
‹Š‡”‹–ƒ…‡ Ž‹‡Ž› „‡ •‹‹Žƒ” ˆ‘” ƒŽŽ
‘”‰ƒ‹••ˆ”‘–Š‹•’Žƒ‡–ǫ
d) ‘—Ž† –Š‡ ‡…Šƒ‹•• ‘ˆ ‰‡‡–‹…
‹Š‡”‹–ƒ…‡ Ž‹‡Ž› „‡ •‹‹Žƒ” –‘
‘”‰ƒ‹••ˆ”‘‡ƒ”–Šǫ
 •Š‘™ ƒ …‘””‡Žƒ–‹‘ ™‹–Š ‘”‰ƒ‹•ƒŽ
”‹…—•‡†–‘†‡†—…‡–Š‡•–”—…–—”‡‘ˆǤ

1.7a)‹•––Š‡†‡ˆ‹‹‰…Šƒ”ƒ…–‡”‹•–‹…•‘ˆ–Š‡
•–”—…–—”‡‘ˆƒ‘Ž‡…—Ž‡Ǥ
b) Š‹…Š ‘ˆ –Š‡•‡ …Šƒ”ƒ…–‡”‹•–‹…• ƒ”‡
‘•–‹’‘”–ƒ––‘”‡’Ž‹…ƒ–‹‘ǫ
c) Š‹…Š …Šƒ”ƒ…–‡”‹•–‹…• ƒ”‡ ‘•–
‹’‘”–ƒ––‘–Š‡‡–”ƒŽ‘‰ƒǫ

1.8‡ˆ‡”–‘ƒ„Ž‡ͳǤͳ
a) Šƒ– ‹• –Š‡ ”‡Žƒ–‹‘•Š‹’ „‡–™‡‡
 …‘–‡– ‘ˆ ƒ ‰‡‘‡ǡ —„‡” ‘ˆ
‰‡‡•ǡ ‰‡‡ †‡•‹–›ǡ ƒ† …Š”‘‘•‘‡
—„‡”ǫ
b)  Šƒ– ˆ‡ƒ–—”‡ ‘ˆ ‰‡‘‡• ‡š’Žƒ‹•
–Š‡…Ǧ˜ƒŽ—‡’ƒ”ƒ†‘šǫ
c)  ‘ ƒ› ‘ˆ –Š‡ —„‡”• ‹ ƒ„Ž‡ ͳǤͳ
1.3 – ‹• ”‡Žƒ–‹˜‡Ž› ‡ƒ•› –‘ ‡š–”ƒ…–  ƒ† …‘’Ž‡š‹–›ǫ
’”‘–‡‹ ˆ”‘ …‡ŽŽ•Ǣ „‹‘…Š‡‹•–• Šƒ† „‡‡ 
†‘‹‰ –Š‹• •‹…‡ ƒ– Ž‡ƒ•– –Š‡ ͳͺͲͲǯ•Ǥ  Š› 1.9 a) ‹•– –Š‡ …Šƒ”ƒ…–‡”‹•–‹…• ‘ˆ ƒ ‹†‡ƒŽ
–Š‡ †‹† ‡”•Š‡› ƒ† Šƒ•‡ ‡‡† –‘ —•‡ ‘†‡Ž‘”‰ƒ‹•Ǥ
”ƒ†‹‘ƒ…–‹˜‹–› –‘ Žƒ„‡Ž  ƒ† ’”‘–‡‹• ‹ b) Š‹…Š ‘†‡Ž ‘”‰ƒ‹• …ƒ „‡ —•‡†
–Š‡‹”‡š’‡”‹‡–•ǫ ‘•–‡ˆˆ‹…‹‡–Ž›–‘‹†‡–‹ˆ›‰‡‡•”‡Žƒ–‡†
 –‘ǣ
1.4  ‘’ƒ”‡ ƒ–•‘ ƒ† ”‹…ǯ• †‹•…‘˜‡”› 
™‹–Š ˜‡”›ǡ ƒ…‡‘† ƒ† …ƒ”–›ǯ•  ‹Ȍ ‡›‡†‡˜‡Ž‘’‡–
†‹•…‘˜‡”›Ǥ  ‹‹Ȍ •‡Ž‡–ƒŽ†‡˜‡Ž‘’‡–
a) Šƒ– †‹† ‡ƒ…Š †‹•…‘˜‡”ǡ ƒ† ™Šƒ–  ‹‹‹Ȍ ’Š‘–‘•›–Š‡•‹•
™ƒ• –Š‡ ‹’ƒ…– ‘ˆ –Š‡•‡ †‹•…‘˜‡”‹‡• ‘  ‹‹‹Ȍ …‡ŽŽ†‹˜‹•‹‘
„‹‘Ž‘‰›ǫ  ‹˜Ȍ …‡ŽŽ†‹ˆˆ‡”‡–‹ƒ–‹‘
b) ‘™ †‹† ƒ–•‘ ƒ† ”‹…ǯ•  ˜Ȍ …ƒ…‡”
ƒ’’”‘ƒ…Š ‰‡‡”ƒŽŽ› †‹ˆˆ‡” ˆ”‘ ˜‡”›ǡ  
ƒ…‡‘†ƒ†…ƒ”–›ǯ•ǫ 1.10‡ˆ‡”–‘ ‹‰—”‡ͳǤͺ
c)”‹‡ˆŽ›”‡•‡ƒ”…Š‘•ƒŽ‹† ”ƒŽ‹‘ a)†‡–‹ˆ›–Š‡’ƒ”–‘ˆ–Š‡‘Ž‡…—Ž‡
–Š‡‹–‡”‡–ǤŠ›‹•Š‡”…‘–”‹„—–‹‘–‘ –Šƒ– ™‘—Ž† „‡ ”ƒ†‹‘ƒ…–‹˜‡Ž› Žƒ„‡Ž‡† ‹
–Š‡•–”—…–—”‡‘ˆ…‘–”‘˜‡”•‹ƒŽǫ –Š‡ƒ‡”—•‡†„›‡”•Š‡›ƬŠƒ•‡
 b)Š‡Ž‹…‡•–Šƒ–ƒ”‡”‹…Š‹
Ǧ„ƒ•‡
1.5–ƒ”–‹‰™‹–Š‹…‡ƒ†ƒ†•–”ƒ‹•‘ˆ ’ƒ‹”• ƒ”‡ Šƒ”†‡” –‘ •‡’ƒ”ƒ–‡ ȋ‡Ǥ‰Ǥ „›
S. pneumoniaeǡ ™Šƒ– ‡š’‡”‹‡–• ‹ Š‡ƒ–‹‰Ȍ–Šƒ–Ǧ”‹…ŠŠ‡Ž‹…‡•ǤŠ›ǫ



Chapter 2 CHROMOSOMES, MITOSIS, AND MEIOSIS 
 

Figure 2.1 Moving

chromosomes (blue)
towards the poles at
anaphase requires
many proteins (red),
all of which interact
with microtubules
  (green).

Chromosomes contain genetic information.  We often take this fact for 
granted,  but  just  over  a  century  ago,  even  the  best  biologists  in  the 
world  were  uncertain  of  the  function  of  these  rod‐shaped  structures. 
We  now  know  that  most  chromosomes  contain  a  single  molecule  of 
double‐stranded  DNA  that  is  complexed  with  proteins.    This 
arrangement  allows  very  long  DNA  molecules  to  be  compacted  into  a 
small volume that can more easily be moved during mitosis and meiosis 
(Fig  2.1).  The  compact  structure  also  makes  it  easier  for  pairs  of 
chromosomes to align with each other during meiosis.  Finally, we shall 
see that chromosomal structure can affect whether genes are active or 
silent. 

CHROMOSOMES MAY BE LOOSE OR COMPACT 
If  stretched  to  its  full  length,  the  DNA  molecule  of  the  largest  human 
chromosome  would  be  85mm.      Yet  during  mitosis  and  meiosis,  this 
DNA  molecule  is  compacted  into  a  chromosome  approximately  5µm 
long.  Although this compaction makes it easier to transport DNA within 
a  dividing  cell,  it  also  makes  DNA  less  accessible  for  other  cellular 
functions  such  as  DNA  synthesis  and  transcription.      Thus, 
chromosomes  vary  in  how  tightly  DNA  is  packaged,  depending  on  the 
stage  of  the  cell  cycle  and  also  depending  on  the  level  of  gene  activity 
required in any particular region of the chromosome.   

There  are  several  different  levels  of  structural  organization  in 

eukaryotic chromosomes, with each successive level contributing to the 
further  compaction  of  DNA  (Fig.  2.2).    For  more  loosely  compacted 
DNA,  only  the  first  few  levels  of  organization  may  apply.    Each  level 
involves  a  specific  set  of  proteins  that  associate  with  the  DNA  to 
C h a p t e r   2  | 2­2  

compact it.  First, proteins called the core histones act as spool around 
which  DNA  is coiled twice to form a structure called the  nucleosome.   
Nucleosomes  are  formed  at  regular  intervals  along  the  DNA  strand, 
giving the molecule the appearance of “beads on a string”.  At the next 
level of organization, histone H1 helps to compact the DNA strand and 
its nucleosomes into a 30nm fibre.  Subsequent levels of organization 
involve the addition of scaffold proteins that wind the 30nm fibre into 
coils, which are in turn wound around other scaffold proteins.  

Figure 2.2 Successive  

stages of chromosome
compaction depend on Chromosomes  stain  very  intensely  with  some  types  of  dyes,  which  is 
the introduction of how they got their name (chromosome means “colored body”).  Certain 
additional proteins. dyes  stain  some  regions  within  a  chromosome  more  intensely  that 
others,  giving  some  chromosomes  a  banded  appearance.  The  material 
that  makes  up  chromosomes,  which  we  now  know  to  be  proteins  and 
DNA,  is  called  chromatin.    There  are  two  general  types  of  chromatin. 
Euchromatin is more loosely packed, and tends to contain more genes 
that are being transcribed, as compared to the more densely compacted 
heterochromatin  which  is  rich  in  short,  repetitive  sequences  called 
microsatellites.  

Chromosomes  also  contain  other  distinctive  features  such  as 

centromeres  and  telomeres.    Both  of  these  are  heterochromatic.    In 
most  cases,  each  chromosome  contains  one  centromere.    These 
Figure 2.3 A pair of sequences are bound by centromeric proteins that link the centromere 
metacentric to  microtubules  that  transport  chromosomes  during  cell  division.   
chromosomes. The Under  the  microscope,  centromeres  can  sometimes  appear  as 
arrow shows a constrictions in the body of the chromosome (Fig. 2.3). If a centromere 
centromeric region. is  located  near  the  middle  of  a  chromosome,  it  is  said  to  be 
metacentric, while an acrocentric centromere is closer to one end of a 
chromosome,  and  a  telocentric  chromsome  is  at  the  very  end.    More 
rarely,  in  a  holocentric  centromere,  no  single  centromere  can  be 
defined and the entire chromsome acts as the centromere.  Telomeres 
are repetitive sequences near the ends of linear chromosomes, and are 
important  in  maintaining  the  length  of  the  chromosomes  during 
replication,  and  protecting  the  ends  of  the  chromosomes  from 
alterations. 

  

 
 C h a p t e r   2  | 2­3  

It  is  useful  to  describe  the  similarity  between  chromosomes  using 
Figure 2.4 appropriate terminology (Fig 2.4). Homologous chromosomes are typically 
Relationships between pairs  of  similar,  but  non‐identical,  chromosomes  in  which  one  member  of 
chromosomes and the pair comes from the male parent, and the other comes from the female 
chromatids. parent.    Homologs  contain  the  same  genes  but  not  necessarily  the  same 
alleles.   Non­homologous chromosomes contain different sets of genes, and 
may  or  may  not  be  distinguishable  based  on  cytological  features  such  as 
length and centromere position.  Within a chromosomes that has undergone 
replication, there are sister chromatids, which are physically connected to 
each other at the centromere and remain joined until cell division.  Because 
a  pair  of  sister  chromatids  is  produced  by  the  replication  of  a  single  DNA 
molecule, their sequences are essentially identical.  On the other hand, non­
sister chromatids come from two separate, but homologous chromosomes, 
and therefore usually contain the same genes in the same order, but do not 
necessarily have identical DNA sequences. 

Figure 2.5
Top: FISH (Fluorescence in situ
hybridization) labeling of all 24
different human chromosomes
(1 - 22, X, and Y) in a fibroblast
nucleus, each with a different
combination of in total seven
fluorochromes.
Bottom: False color
representation of all
chromosome territories visible
in this mid-section after
computer classification.  
  

  
C h a p t e r   2  | 2­4  

MITOSIS 
Cell  division  is  essential  to  asexual  reproduction  and  the  development 
of multicellular organisms.  Accordingly, the primary function of mitosis 
is  to  ensure  that  each  daughter  cell  inherits  identical  genetic  material, 
i.e.  exactly  one  copy  of  each  chromosome.    To  make  this  happen, 
replicated  chromosomes  condense  (prophase),  and  are  positioned 
near  the  middle  of  the  dividing  cell  (metaphase),  and  then  one  sister 
chromatid from each chromosome migrates towards opposite poles of 
the  dividing  cell  (anaphase),  until  the  identical  sets  of  chromosomes 
are  completely  separated  from  each  other  within  the  newly  formed 
nuclei of each daughter cell (telophase) (see Figs. 2.5‐2.7 for diagrams 
of  the  process).      This  is  followed  by  the  completion  of  the  division  of 
the  cytoplasm  (cytokinesis).  The  movement  of  chromosomes  is  aided 
by microtubules that attach to the chromosomes at centromere.    

Figure 2.6 Mitosis in

arabidopsis showing
fluorescently labeled
chromosomes (blue)
and microtubules
(green) at metaphase,
anaphase and  
telophase (from left to
right).  

MEIOSIS 
Meiosis, like mitosis, is also a necessary part of cell division.  However, 
in  meiosis  not  only  do  sister  chromatids  separate  from  each  other, 
homologous  chromosomes  also  separate  from  each  other.    This  extra, 
reductional step of meiosis is essential to sexual reproduction.  Without 
meiosis, the chromosome number would double in each generation of a 
species and would quickly become too large to be viable.     

Meiosis is divided into two stages designated by the roman numerals I 
and II.  Meiosis I is called a reductional division, because it reduces the 
number  of  chromosomes  inherited  by  each  of  the  daughter  cells.  
Meiosis  I  is  further  divided  into  Prophase  I,  Metaphase  I,  Anaphase  I, 
and Telophase I, which are roughly similar to the corresponding stages 
of  mitosis,  except  that  in  Prophase  I  and  Metaphase  I,  homologous 
chromosomes  pair  with  each  other  in  transient  structures  called 
bivalents  (Figs.  2.7,  2.8).    This  is  an  important  difference  between 
mitosis  and  meiosis,  because  it  affects  the  segregation  of  alleles,  and 
also  allows  for  recombination  to  occur  through  crossing‐over,  as 
described later in the course.  During Anaphase I, one member of each 
pair  of  homologous  chromosomes  migrates  into  a  daughter  cell.  
Meiosis II is essentially the same as mitosis, with one sister chromatid 
from  each  chromosome  separating  to  produce  two  identical  cells.  
Because  Meiosis  II,  like  mitosis,  results  in  products  that  contain 
identical sequences, Meiosis II is called an equational division. 
  

 
 C h a p t e r   2  | 2­5  

Figure 2.7 Mitosis and meiosis. Note the similarities and differences between metaphase in
mitosis and metaphase I and II of meiosis.
 

Figure 2.8 Meiosis in Arabidopsis (n=5). Panels A-C show different stages of prophase I, each with an
increasing degree of chromosome condensation. Subsequent phases are shown: metaphase I (D), telophase I
(E), metaphase II (F), anaphase II (G), and telophase II (H).
  
C h a p t e r   2  | 2­6  

 
 

THE CELL CYCLE AND CHANGES IN DNA CONTENT 
The life cycle of an eukaryotic cell can generally be divided into at least 
four stages (Fig. 2.9).  When a cell is produced through fertilization or 
cell  division,  there  is  usually  a  lag  before  it  undergoes  DNA  synthesis.  
This lag period is called Gap 1 (G1), and ends with the onset of the DNA 
synthesis  (S)  phase,  during  which  each  chromosome  is  replicated.  
Following  replication,  there  may  be  another  lag,  called  Gap  2  (G2), 
Figure 2.9 A typical before mitosis (M).  Cells undergoing meiosis do not usually have a G2 
eukaryotic cell cycle.
phase.      Interphase  is  as  term  used  to  describe  all  phases  of  the  cell 
cycle  excluding  mitosis  or  meiosis.  A  typical  cell  cycle  is  shown  in  Fig. 
2.9.    My  variants  of  this  generalized  cell  cycle  also  exist.    Some  cells 
never  leave  G1  phase,  and  are  said  to  enter  a  permanent,  non‐dividing 
stage called G0.  On the other hand, some cells undergo many rounds of 
DNA  synthesis  (S)  without  any  mitosis  or  cell  division.    These 
endoreduplicated  cells  are  described  later  in  this  chapter.  
Understanding the control of the cell cycle is an active area of research, 
particularly  because  of  the  relationship  between  cell  division  and 
cancer. 

The  amount  of  DNA  within  a  cell  changes  following  each  of  the 
following events: fertilization, DNA synthesis, mitosis, and meiosis (Fig 
2.10).    We  use  “c”  to  represent  the  DNA  content  in  a  cell,  and  “n”  to 
represent  the  number  of  complete  sets  of  chromosomes.    In  a  gamete 
(i.e.  sperm  or  egg),  the  amount  of  DNA  is  1c,  and  the  number  of 
chromosomes  is  1n.    Upon  fertilization,  both  the  DNA  content  and  the 
number of chromosomes doubles to 2c and 2n, respectively.   Following 
DNA  synthesis,  the  DNA  content  doubles  again  to  4c,  but  each  pair  of 
sister  chromatids  is  still  counted  as  a  single  chromosome,  so  the 
number  of  chromosomes  remains  unchanged  at  2n.    If  the  cell 
undergoes mitosis, each daughter cell will be 2c and 2n, because it will 
receive  half  of  the  DNA,  and  one  of  each  pair  of  sister  chromatids.    In 
  

 
 C h a p t e r   2  | 2­7  

contrast, the cells that are produced from the meiosis of a 2n, 4c cell are 
1c  and  1n,  since  each  pair  of  sister  chromatids,  and  each  pair  of 
homologous chromosomes divides during meiosis. 

Figure 2.10
Changes in DNA
and chromosome
content during
the cell cycle.
For simplicity,
nuclear
membranes are
not shown, and
all chromosomes
are represented
in a similar stage
of condensation.  

  
C h a p t e r   2  | 2­8  

KARYOTYPES SHOW CHROMOSOME NUMBER AND STRUCTURE 
Each  eukaryotic  species  has  its  total  nuclear  genome  divided  among  a 
number  of  chromosomes  that  is  characteristic  of  that  species.    For 
example, a haploid human nucleus (i.e. sperm or egg) normally has 23 
chromosomes  (n=23),  and  a  diploid  human  nucleus  has  23  pairs  of 
chromosomes  (2n=46).    Various  stains  and  fluorescent  dyes  produce 
characteristic banding patterns on some chromosomes.  This can make 
it  easier  to  identify  specific  chromosomes.    The  number  of 
chromosomes varies between species (see Table 1.1), but there appears 
to  be  very  little  correlation  between  chromosome  number  and  either 
the complexity of an organism or its total amount genomic DNA.   

Figure 2.11 Karyotype

of a normal human
male
 
A karyotype shows the complete set of chromosomes of an individual 
(Fig.  2.11).  Analysis  of  karyotypes  can  identify  of  chromosomal 
abnormalities,  including  aneuploidy,  which  is  the  addition  or 
subtraction  of  a  chromosome  from  a  pair  of  homologs.    More 
specifically,  the  absence  of  one  member  of  a  pair  of  homologous 
chromosomes  is  called  monosomy.    On  the  other  hand,  in  a  trisomy, 
there are three, rather than two homologs of a particular chromosome.   
Different types of aneuploidy are sometimes represented symbolically; 
if  2n symbolizes the normal number of chromosomes in a cell, then 2n­
1 indicates monosomy and 2n+1 represents trisomy.   

The most familiar human aneuploidy is trisomy‐21 (i.e. three copies of 
chromosome 21), which is one cause of Down’s syndrome.  Most (but 
not  all)  other  human  aneuploidies  are  lethal  at  an  early  stage  of 
  

 
 C h a p t e r   2  | 2­9  

embryonic development.  Note that aneuploidy usually affects only one 
set  of  homologs  within  a  karyotype,  and  is  therefore  distinct  from 
polyploidy,  in  which  the  entire  karyotype  is  duplicated  (see  below).  
Aneuploidy  is  almost  always  deleterious,  whereas  polyploidy  appears 
to be beneficial in some organisms, particularly some species of plants. 

  Figure 2.12
  Structural abberations
in chromosomes.
 

Structural defects in chromosomes are another type of abnormality that 
can  be  detected  in  karyotypes  (Fig  2.12).    These  defects  include 
deletions, duplications, and inversions, which all involve changes in a 
segment  of  a  single  chromosome.  Insertions  and  translocations 
involve two non‐homologous chromosomes.  In an insertion, DNA from 
one  chromosome  is  unidirectional,  while  in  translocation,  the  transfer 
of  chromosomal  segments  is  bidirectional.      Structural  defects  affect 
only  part  of  a  chromosome,  and  so  tend  to  be  less  harmful  than 
aneuploidy.  In  fact,  there  are  many  examples  of  ancient  chromosomal 
rearrangements  in  the  genomes  of  species  including  our  own.  
Duplications  of  some  small  chromosomal  segments,  in  particular,  may 
have  some  evolutionary  advantage  by  providing  extra  copies  of  some 
genes, which can then evolve in new ways. 

Chromosomal  abnormalities  arise  in  many  different  ways.    Many  of 

these  can  be  traced  to  rare  errors  in  natural  cellular  processes.  Non­
disjunction  is  the  failure  of  at  least  one  pair  of  chromosomes  or 
chromatids  to  separate  during  mitosis  or  meiosis.    Chromosome 
breakage  also  occurs  infrequently  as  the  result  of  physical  damage 
(such as radiation), movement of some types of transposons, and other 
factors.    During  the  repair  of  a  broken  chromosome,  deletions, 
insertions, translocations and even inversions can be introduced.     

  
C h a p t e r   2  | 2­10  

POLYPLOIDY 
Humans,  like  most  animals  and  all  of  the  eukaryotic  genetic  model 
organisms in wide use, are diploid.  This means that most of their cells 
have  two  homologous  copies  of  each  chromosome.    In  contrast,  many 
plant  species  and  even  a  few  animal  species  are  polyploids.    This 
means there are more than two homologs of each chromosome in each  
cell.   

When describing polyploids, we use the letter “x” to define the level of 
ploidy.    A  diploid  is  2x,  because  there  are  two  basic  sets  of 
chromosomes, and a tetraploid is 4x, because it contains four copies of 
each chromosome.  For clarity when discussing polyploids,  geneticists 
will  often  combine  the  “x”  notation  with  the  “n”  notation  already 
defined  previously  in  this  chapter.    Thus  for  both  diploids  and 
polyploids, “n” is the number of chromosomes in a gamete, and “2n” is 
the  number  of  chromosomes  following  fertilization.    For  a  diploid, 
therefore, n=x, and 2n=2x.  For a tetraploid, n=2x, and 2n=4x.  

Like diploids (2n=2x), stable polyploids generally have an even number 
of  copies  of  each  chromosome:  tetraploid  (2n=4x),  hexaploid  (2n=6x), 
and so on.  The reason for this is clear from a consideration of meiosis.  
Remembering  that  the  purpose  of  meiosis  is  to  reduce  the  sum  of  the 
genetic material by half, meiosis can equally divide an even number of 
chromosome  sets,  but  not  an  odd  number.    Thus,  polyploids  with  an 
uneven  number  of  chromosomes  (e.g.  triploids,  2n=3x)  tend  to  be 
sterile, even if they are otherwise healthy.  The mechanism of meiosis in 
stable  polyploids  is  essentially  the  same  as  in  diploids:  during 
metaphase  I,  homologous  chromosomes  pair  with  each  other.  
Depending on the species, all of the homologs may be aligned together 
at  metaphase,  or  in  multiple  separate  pairs.    For  example,  in  a 
tetraploid,  some  species  may  form  tetravalents  in  which  the  four 
homologs  from  each  chromosome  align  together,  or  alternatively,  two 
pairs  of  homologs  may  form  two  bivalents.  Note  that  because  that 
mitosis  does  not  involve  any  pairing  of  homologous  chromosomes, 
mitosis  is  equally  effective  in  diploids,  even‐number  polyploids,  and 
odd‐number polyploids.       

Triploidy  is  used  in  the  production  of  seedless  fruits,  such  as 
watermelon,  grapes  and  bananas.    All  of  the  tissues  of  these  fruit  are 
triploid.  Because almost all of the cells of the plant, including its fruit, 
are produced through mitosis, the uneven number of chromosome sets 
does not affect their development.  However, cells that contribute to the 
production of gametes are produced through meiosis, and because the 
triploids  are  unable  to  complete  normal  meioses,  their  gametes  fail  to 
develop,  so  no  zygotes  are  formed,  and  the  seeds  (which  normally 
contain embryos that develop from the zygotes) are aborted.  

  

 
 C h a p t e r   2  | 2­11  

If  triploids  cannot  make  seeds,  how  do  we 

obtain  enough  triploid  individuals  for 
cultivation?  The answer depends on the plant 
species  involved.  In  some  cases,  such  as 
banana,  it  is  possible  to  propagate  the  plant 
asexually;  new  progeny  can  simply  be  grown 
from  cuttings  from  a  triploid  plant.    On  the 
other hand, seeds for seedless watermelon are 
produced  sexually:  a  tetraploid  watermelon  Figure 2.13. Part of a
plant  is  crossed  with  a  diploid  watermelon  triploid watermelon,
plant.  Both the tetraploid and the diploid are  showing white, aborted
fully  fertile,  and  produce  gametes  with  two  seeds within the flesh
(1n=2x) or one (1n=1x) sets of chromosomes, 
respectively.    These  gametes  fuse  to  produce  a  zygote  (2n=3x),  that  is 
able to develop normally into an adult plant through multiple rounds of 
mitosis, but is unable to compete normal meiosis or produce seeds. 

Figure 2.14.
Endoreduplicated
chromosomes from an
insect salivary gland.
The banding pattern is
produced with
fluorescent labels.  

ENDOREDUPLICATION 
Endoreduplication,  also  known  as  endopolyploidy,  is  a  special  type 
of  tissue‐specific  genome  amplification  that  occurs  in  many  types  of 
plant  cells  and  in  specialized  cells  of  some  animals  including  humans.  
Endoreduplication  does  not  affect  the  germline  or  gametes,  so  species 
with  endopolyploidy  are  not  considered  polyploids.    Endopolyploidy 

  
C h a p t e r   2  | 2­12  

occurs  when  a  cell  undergoes  multiple  rounds  of  DNA  synthesis  (S‐
phase) without any mitosis.  This produces multiple chromatids of each 
chromosome.    Endopolyploidy  seems  to  be  associated  with  cells  that 
are  metabolically  very  active,  and  produce  a  lot  of  enzymes  and  other 
proteins  in  a  short  amount  of  time.    The  highly  endoreduplicated 
salivary  gland  chromosomes  of  D.  melanogaster  have  been  useful 
research  models  in  genetics,  since  their  relatively  large  size  makes 
them easy to study under the microscope.  

GENE BALANCE 
Why do trisomies, duplications, and other chromosomal abnormalities 
that  increase  gene  copy  number  sometimes  have  a  negative  effect  on 
the  normal  development  or  physiology  of  an  organism?    This  is 
particularly  intriguing  because  in  many  species,  aneuploidy  is 
detrimental  or  lethal,  while  polyploidy  is  tolerated  or  even  beneficial.  
The answer is probably related to the concept of gene balance, which  
can  be  summarized  as  follows:  genes,  and  the  proteins  they  produce, 
have evolved to be part of complex metabolic and regulatory networks.  
Some  of  these  networks  function  best  when  certain  enzymes  and 
regulators  are  present  in  specific  ratios  to  each  other.    Increasing  the 
gene  copy  number  for  just  one  part  of  the  network  may  throw  the 
network  out  of  balance,  leading  to  increases  or  decreases  of  certain 
metabolites,  which  may  be  toxic  in  high  concentrations  or  which  may 
be  limiting  in  other  important  processes  in  the  cell.    The  activity  of 
genes  and  metabolic  networks  is  regulated  in  many  different  ways 
besides changes in gene copy number, so duplication of just a few genes 
will  usually  not  be  harmful.    However,  trisomy  and  large  segmental 
duplications  of  chromosomes  affect  the  dosage  of  so  many  genes  that 
cellular networks are unable to adjust to the changes. 

ORGANELLAR GENOMES 
Chromosomes  also  exist  outside  of  the  nucleus,  within  both  the 
chloroplast and mitochondria.  These organelles are likely the remnants 
of  a  prokaryotic  endosymbionts  that  entered  the  cytoplasm  of  ancient 
progenitors  of  today’s  eukaryotes.    These  endosymbionts  had  their 
own,  circular  chromosomes,  like  most  bacteria  that  exist  today.  
Likewise,  chloroplasts  and  mitochondria  also  have  circular 
chromosomes  that  behave  more  like  bacterial  chromosomes  than 
eukaryotic chromosomes, i.e. these organellar genomes do not undergo 
mitosis  or  meiosis.    Organellar  genomes  are  also  often  present  in 
multiple copies within each organelle, and in most species are inherited 
maternally. 

  

 
 C h a p t e r   2  | 2­13  

_______________________________________________________________________________________ 

SUMMARY 
• Chromosomes  are  complex  and  dynamic  structures  consisting  of  DNA  and  proteins 
(chromatin). 
 
• The  degree  of  chromatin  compaction  varies  between  heterochromatic  and  euchromatic 
regions and between stages of the cell cycle. 
 
• Some chromosomes can be distinguished cytologicaly based on their length, centromere 
position,  and  banding  patterns  when  stained  dyes  or  labelled  with  sequence‐specific 
probes. 
 
• Homologous chromosomes contain the same genes, but not necessarily the same alleles.  
Sister chromatids usually contain the same genes and the same alleles. 
 
• Mitosis reduces the c‐number, but not the n‐number.  Meiosis reduces both c and n. 
 
• Homologous chromosomes associate with each other during meiosis, but not mitosis. 
 
• Several types of structural defects in chromosomes occur naturally, and can affect cellular 
function and even evolution.  
 
• Aneuploidy results from the addition or subtraction of one or more chromosomes from a 
group of homologs, and is usually deleterious to the cell. 
 
• Polyploidy is the presence of more than two complete sets of chromosomes in a genome.  
Even‐numbered  multiple  sets  of  chromosomes  can  be  stably  inherited  in  some  species, 
especially plants. 
 
• Endopolyploidy is tissue‐specific type of polyploidy observed in some species, including 
diploids. 
 
• Both aneuploidy and structural defects such as duplications can affect gene balance. 
 
• Organelles  also  contain  chromosomes,  but  these  are  much  more  like  prokaryotic 
chromosomes than the nuclear chromosomes of eukaryotes. 
 

  
C h a p t e r   2  | 2­14  

KEY TERMS 
chromosome  mitosis  c 
core histones  prophase  karyotype 
nucleosome  metaphase  aneuploidy 
30nm fiber  anaphase  monsomic 
histone H1  telophase  trisomic 
scaffold proteins  prophase  Down’s syndrome 
heterochromatin  meiosis  deletion 
euchromatin  prophase (I, II)  duplication 
microsatellite  metaphase (I, II)  insertion 
chromatid  anaphase (I, II)  inversion 
centromere  telophase (I, II)  translocation 
metacentric  cytokinesis  non‐disjunction 
acrocentric  bivalent  chromosome breakage 
telocentric  reductional division  polyploid 
holocentric  equational division  x 
telomere  G 1  tetravalent 
homolog  G 2  endoreduplication 
non‐homologous  S  endopolyploidy 
chromatid  M  gene balance 
sister chromatid  G 0  cellular network 
non‐sister chromatid  interphase  endosymbiont 
interphase  n  organellarchromo
 

_______________________________________________________________________________________ 

STUDY QUESTIONS 
2.1 Define chromatin. What is the difference  2.4 
between DNA, chromatin and  a)  How  many  centromeres  does  a  typical 
chromosomes?  chromosome have?  
  b)  What  would  happen  if  there  was  more 
2.2  Species  A  has  n=4  chromosomes  and  than one centromere per chromosome?  
Species B has n=6 chromosomes. Can you  c)  What  if  a  chromosome  had  zero 
tell from this information which species has  centromeres? 
more DNA? Can you tell which   
species has more genes?  2.5 For a diploid with 2n=16 chromosomes, 
  how  many  chromosomes  and  chromatids 
2.3  The  answer  to  question  2  implies  that  are  per  cell  present  in  the  gamete,  and 
not all DNA within a chromosome encodes  zygote  and  immediately  following  G1,  S,  G2, 
genes.  Can  you  name  any  examples  of  mitosis, and meiosis?   
chromosomal regions that contain   
relatively few genes?  2.6  Bread  wheat  (Triticum  aestivum)  is  a 
   hexaploid.  Using  the  nomenclature 
  presented  in  class,  an  egg  cell  of  wheat  has 
 
  

 
 

n=21  chromosomes.  How  many   

chromosomes in a zygote of bread wheat?  2.9  For  a  diploid  organism  with  2n=4 
  chromosomes,  draw  a  diagram  of  all  of  the 
2.7 For a given gene:  possible  configurations  of  chromosomes 
a)  What  is  the  maximum  number  of  alleles  during  normal  anaphase  I,  with  the 
that  can  exist  in  a  2n  cell  of  a  given  diploid  maternally  and  paternally  derived 
individual?  chromosomes labelled.      
b)  What  is  the  maximum  number  of  alleles   
that  can  exist  in  a  1n  cell  of  a  tetraploid  2.10  For  a  triploid  organism  with  2n=3x=6 
individual?  chromosomes,  draw  a  diagram  of  all  of  the 
c)  What  is  the  maximum  number  of  alleles  possible  configurations  of  chromosomes  at 
that  can  exist  in  a  2n  cell  of  a  tetraploid  anaphase  I  (it  is  not  necessary  label 
individual?  maternal and paternal chromosomes).      
d)  What  is  the  maximum  number  of  alleles   
that can exist in a population?  2.11    For  a  tetraploid  organism  with 
  2n=4x=8  chromosomes,  draw  all  of  the 
2.8   possible  configurations  of  chromosomes 
a) Why is aneuploidy more often lethal than  during a normal metaphase.  
polyploidy?     
b)  Which  is  more  likely  to  disrupt  gene 
balance: polyploidy or duplication? 
 

 
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in peas. %.0#&L/,1%(0!51,-%3,!;(%51(<! /.%3,!,-#!&#*/,1%(0-1:0!.#,5##(!<#(#0+!
2-&%9%0%9#0+!/()!CB@6!!J#!(%5!;(%5!,-/,!,-#!&#/0%(!,-/,!9%&#!,-/(!
%(#! /**#*#! %$! /! <#(#! 2/(! .#! :&#0#(,! 1(! /(! 1()1L1)3/*! 10! ,-/,! 9%0,!
#3;/&4%,12! %&</(1090! -/L#! /,! *#/0,! ,5%! 0#,0! %$! -%9%*%<%30!
2-&%9%0%9#06! ! 7%&! %&</(1090! ,-/,! /&#! :&#)%91(/(,*4! )1:*%1)+!032-! /0!
-39/(0! %&! '#()#*E0! :#/0+! 2-&%9%0%9#0! #810,! 1(! :/1&0+! 51,-! %(#!
-%9%*%<!1(-#&1,#)!$&%9!#/2-!:/&#(,6!!!C1:*%1)!2#**0!2/(!,-#&#$%&#!-%*)!
3:!,%!,5%!)1$$#&#(,!/**#*#0!%$!#/2-!<#(#+!51,-!%(#!/**#*#!%(!#/2-!9#9.#&!
%$! /! :/1&! %$! -%9%*%<%30! 2-&%9%0%9#06! ! G$! .%,-! /**#*#0! %$! /! :/&,123*/&!
<#(#! /&#! 1)#(,12/*+! ,-#! 1()1L1)3/*! 10! 0/1)! ,%! .#! 01:1;<715$! $%&! ,-/,!
<#(#6!!Q(!,-#!%,-#&!-/()+!1$!,-#!/**#*#0!/&#!)1$$#&#(,!$&%9!#/2-!%,-#&+!,-#!
<#(%,4:#! 10! 0#,#+1;<715$6! ! G(! 0%9#! 2/0#0+! 032-! /0! 5-#(! )#02&1.1(<!
<#(#0! %(! ,-#! 0#8! 2-&%9%0%9#0! %$! /! -39/(! 9/*#+! 5#! 30#! ,-#! ,#&9!
0#:*;<715$6! ! I-10! 10! .#2/30#! ,-#! 2#**! 2%(,/1(0! %(*4! %(#! U! /()! %(#! V!
2-&%9%0%9#+! 0%! ,-#&#! 10! %(*4! %(#! :%001.*#! /**#*#! $%&! #/2-! <#(#6!!
@*,-%3<-! /! ,4:12/*! )1:*%1)! 1()1L1)3/*! 2/(! -/L#! /,! 9%0,! ,5%! )1$$#&#(,!
/**#*#0! %$! /! :/&,123*/&! <#(#+! 9%&#! ,-/(! ,5%! /**#*#0! 2/(! #810,! 1(! /!
:%:3*/,1%(! %$! 1()1L1)3/*06! ! I-#! 9%0,! 2%99%(! /**#*#! 1(! /! (/,3&/*!
:%:3*/,1%(!10!2/**#)!,-#!.*"'",<=#!/**#*#6!!!!

HAD@IGQBFWGXF!"AIJAAB!@DDADAF!@BC!XWABQIVXAF!
@!0:#21$12!:%01,1%(!%(!/!2-&%9%0%9#!10!2/**#)!/!"1>5$6!!'%0,!%$!,-#!*%21!
<#(#,1210,0!)102300!/&#!%223:1#)!.4!<#(#0+!/()!,-#!,#&90!*%230!/()!<#(#!
/&#!%$,#(!30#)!1(,#&2-/(<#/.*46!!I-#!2%9:*#,#!0#,!%$!/**#*#0!/,!/(4!*%21!
%$!1(,#&#0,!1(!/(!1()1L1)3/*!)#$1(#!1,0!7#&1,<=#6!!I-#!)#,#2,/.*#!#$$#2,!%$!
,-#0#!/**#*#0!%(!,-#!0,&32,3&#!%&!$3(2,1%(!%$!,-/,!1()1L1)3/*!10!2/**#)!1,0!

!
 Q ( # ! D % 2 3 0 !S!8"8!!

=0#&1,<=#6! ! ! I-#! :-#(%,4:#! 0,3)1#)! 1(! /(4! :/&,123*/&! <#(#,12!

#8:#&19#(,!9/4!&/(<#!$&%9!019:*#+!L101.*#!,&/1,0!032-!/0!-/1&!2%*%&+!,%!
9%&#!2%9:*#8!:-#(%,4:#0!1(2*3)1(<!)10#/0#!0302#:,1.1*1,4!%&!.#-/L1%&6!!!

Figure 3.3
Relationship between
genotype and
phenotype for a
dominant allele.

!
D#,! 30! &#,3&(! ,%! /(! #8/9:*#! %$! /! 019:*#! :-#(%,4:#M! $*%5#&! 2%*%&! 1(!
'#()#*E0! :#/06! J#! -/L#! /*&#/)4! 0/1)! ,-/,! %(#! /**#*#! :&%)32#0! :3&:*#!
$*%5#&0+!5-1*#!,-#!%,-#&!/**#*#!:&%)32#0!5-1,#!$*%5#&0!Y71<3&#!O6OZ6!!"3,!
5-/,! )%#0! ,-10! ,#**! 30! /.%3,! ,-#! $*%5#&! 2%*%&! %$! /(! 1()1L1)3/*! ,-/,! -/0!
%(#! :3&:*#! /**#*#! /()! %(#! 5-1,#! /**#*#+! 1(! %,-#&! 5%&)0+! 5-/,! 10! ,-#!
=0#&1,<=#!%$!/(!1()1L1)3/*!5-%0#!<#(%,4:#!10!-#,#&%K4<%30?!J#!;(%5!
$&%9! #8:#&19#(,/*! )/,/! ,-/,! 1()1L1)3/*0! -#,#&%K4<%30! $%&! ,-#! :3&:*#!
/()! 5-1,#! /**#*#0! %$! ,-#! $*%5#&! 2%*%&! <#(#! -/L#! :3&:*#! $*%5#&06! ! ! ! I-#!
/**#*#!/00%21/,#)!51,-!:3&:*#!2%*%&!10!,-#&#$%&#!0/1)!,%!.#!'1:*&!&,!,%!
,-#! /**#*#! ,-/,! :&%)32#0! ,-#! 5-1,#! 2%*%&6! ! I-#! %,-#&! /**#*#+! 5-%0#!
:-#(%,4:#! 10! 9/0;#)! .4! /! )%91(/(,! /**#*#! 1(! /! -#,#&%K4<%,#+! 10!2/**#)!
+#>#$$*?#6! ! ! Q$,#(+! /! )%91(/(,! /**#*#! 51**! .#! &#:&#0#(,#)! .4! /! 2/:1,/*!
*#,,#&!Y#6<6!!Z!5-1*#!/!&#2#001L#!/**#*#!51**!.#!&#:&#0#(,#)!1(!*%5#&!2/0#!
Y#6<6!"Z6!!W%5#L#&+!9/(4!)1$$#&#(,!040,#90!%$!<#(#,12!049.%*0!/&#!1(!30#!
YI/.*#! O6[Z+! 0%! 1,! 10! 19:%&,/(,! ,%! 3()#&0,/()! ,-#! )1$$#&#(,! ,4:#0! %$!
(%,/,1%(!/()!,%!30#!,-#9!2%(010,#(,*46!!!

Table 3.1 Examples of symbols used to represent alleles, and their dominance relationships
#8/9:*#0!%$! ,4:12/*!1(,#&:&#,/,1%(
<#(%,4:#0!%$!
-#,#&%K4<%,#0!

!"! R::#&2/0#!*#,,#&Y0Z 9%0,!%$,#(!&#:&#0#(,0!)%91(/(,!/**#*#+!/()!

*%5#&2/0#!*#,,#&Y0Z!1()12/,#0!&#2#001L#!/**#*#6!!!

"#"$%&! D%5#&2/0#!*#,,#&0!51,-!\!03:#&02&1:,!&#:&#0#(,!51*)=,4:#!/**#*#+!
5-12-!10!%$,#(!)%91(/(,6!!@!*%5#&!2/0#!*#,,#&!51,-!%,-#&!
! 03:#&02&1:,0!Y%&!51,-!(%!03:#&02&1:,0Z!&#:&#0#(,0!%,-#&!/**#*#0+!
5-12-!/&#!%$,#(!&#2#001L#6!

!'!&! I-#!(/9#!%$!,-#!*%230!10!1(!3::#&2/0#!%&!*%5#&!2/0#!*#,,#&0+!51,-!
03.02&1:,0!%&!03:#&02&1:,0!30#)!,%!1()12/,#!)1$$#&#(,!/**#*#06!!
P#(#&/**4+!(#1,-#&!/**#*#!10!2%9:*#,#*4!)%91(/(,+!%&!#*0#!,-#!
)%91(/(2#!&#*/,1%(0-1:0!/&#!3(;(%5(6!!

!
 Q ( # ! D % 2 3 0 !S!8"@!!

"#01)#0! )%91(/(2#! /()! &#2#001L1,4+! %,-#&! &#*/,1%(0-1:0! 2/(! #810,!

.#,5##(! /**#*#06! ! G(! $#:*"'1:*&!&>#! Y/*0%! 2/**#)! *&>1:="#,#!
'1:*&!&>#A!71<3&#!O6]Z+!.%,-!/**#*#0!/$$#2,!,-#!,&/1,!/))1,1L#*4+!/()!,-#!
:-#(%,4:#! %$! ,-#! -#,#&%K4<%,#! 10! 1(,#&9#)1/,#! .#,5##(! #1,-#&! %$! ,-#!
-%9%K4<%,#06! ! 7%&! #8/9:*#+! /**#*#0! %$! $%&! &#)! 2%*%&! 1(! 2/&(/,1%(0! /()!
0%9#!%,-#&!0:#21#0!#8-1.1,!0#91=)%91(/(2#+!0%!,-/,!/!-#,#&%K4<%,#!-/0!
:1(;! :#,/*0+! 5-1*#! /! -%9%K4<%,#! $%&! %(#! /**#*#! -/0! &#)! :#,/*0! /()! ,-#!
%,-#&! -/0! 5-1,#! :#,/*06! ! ! I-30! 1(! 0#91=)%91(/(2#+! ,-#! )%0/<#! %$! ,-#!
/**#*#0!/$$#2,0!,-#!:-#(%,4:#+!#6<6!,-#!:-#(%,4:#!%$!,-#!-#,#&%K4<%,#!10!
/::&%819/,#*4!-/*$!/0!0,&%(<!/0!,-#!:-#(%,4:#!%$!/!-%9%K4<%,#6!!!!!!

Figure 3.4
Relationship between
genotype and
phenotype for semi-
dominant alleles.
!
!

B1"'1:*&!&>#! 10! /(%,-#&! ,4:#! %$! /**#*12! &#*/,1%(0-1:+! 1(! 5-12-! /!

-#,#&%K4<%30! 1()1L1)3/*! #8:&#00#0! ,-#! :-#(%,4:#! %$! .%,-! /**#*#0!
0193*,/(#%30*46!!@(!#8/9:*#!%$!2%=)%91(/(2#!10!$%3()!51,-1(!,-#!@"Q!
.*%%)! <&%3:! %$! -39/(0+! 5-12-! 10! 2%(,&%**#)! .4! /! <#(#! 2/**#)!(! Y71<3&#!
O6^Z6! ! I-#&#! /&#! ,-&##! :%001.*#! /**#*#0! /,! ,-10! *%230M! (!+! ()+! /()! *6!!!
W%9%K4<%30!1()1L1)3/*0!$%&!(!!%&!()!:&%)32#!%(*4!@!%&!"!,4:#!/(,1<#(0+!
&#0:#2,1L#*4+! %(! ,-#! 03&$/2#! %$! ,-#1&! .*%%)! 2#**0+! /()! ,-#&#$%&#! -/L#!
#1,-#&!,4:#!@!%&!,4:#!"!.*%%)6!!W#,#&%K4<%30!(!()!1()1L1)3/*0!-/L#!.%,-!
@! /()! "! /(,1<#(0! %(! ,-#1&! 2#**! 03&$/2#+! /()! 0%! -/L#! ,4:#! @"! .*%%)6!!
B%,12#!,-/,!,-#!-#,#&%K4<%,#!#8:&#00#0!.%,-!/**#*#0!0193*,/(#%30*4+!/()!
10!(%,!0%9#!;1()!%$!(%L#*!1(,#&9#)1/,#!.#,5##(!@!/()!"6!!N%=)%91(/(2#!
10! ,-#&#$%&#! )10,1(2,! $&%9! 0#91=)%91(/(2#+! /*,-%3<-! ,-#4! /&#!
0%9#,19#0!2%($30#)6!!!'/(4!,4:#0!%$!9%*#23*/&!9/&;#&0+!5-12-!5#!51**!
)102300! 1(! /! */,#&! 2-/:,#&+! )10:*/4! /! 2%=)%91(/(,! &#*/,1%(0-1:! /9%(<!
/**#*#06!!G,!10!/*0%!19:%&,/(,!,%!(%,#!,-/,!,-#!,-1&)!/**#*#+!*+!)%#0!(%,!9/;#!
/(4! /(,1<#(0! /()! 10! &#2#001L#! ,%! /**! %,-#&! /**#*#06! ! ! W%9%K4<%30!
&#2#001L#! 1()1L1)3/*0! Y**Z! -/L#! ,4:#! Q! .*%%)6! I-10! 10! /! 30#$3*! &#91()#&!
,-/,! )1$$#&#(,! ,4:#0! %$! )%91(/(2#! &#*/,1%(0-1:0! 2/(! #810,+! #L#(! $%&!
/**#*#0!%$!,-#!0/9#!<#(#6!

!
 Q ( # ! D % 2 3 0 !S!8"C!!

Figure 3.5
Relationship between
genotype and
phenotype for co-
! dominant alleles (IA,
IB), and a recessive
"GQNWA'GN@D!"@FGF!Q7!CQ'GB@BNA! allele.
J#!2/((%,!:&#)12,!5-#,-#&!/(!/**#*#!51**!.#!)%91(/(,!./0#)!019:*4!%(!
,-#!:-#(%,4:#!%$!-%9%K4<%,#0>!5#!930,!/*0%!;(%5!,-#!:-#(%,4:#!%$!/!
-#,#&%K4<%,#6! ! J-/,+! ,-#(+! 2/30#0! )%91(/(2#?! ! I-#&#! /&#! 9/(4!
)1$$#&#(,!.1%2-#912/*!2%()1,1%(0!,-/,!9/4!9/;#!%(#!/**#*#!)%91(/(,!,%!
/(%,-#&+!.3,!%(#!%$!,-#!9%0,!2%99%(!10!0!="1$522*>*#&><6!!'%0,!<#(#0!
:&%)32#! 9%&#! ,-/(! #(%3<-! :&%,#1(! ,%! /22%9:*10-! ,-#1&! (%&9/*!
.1%*%<12/*!$3(2,1%(6!!G$!/(!/**#*#!10!-/:*%03$$121#(,+!,-#(!_30,!%(#!2%:4!%$!
,-/,! /**#*#! :&%)32#0! #(%3<-! :&%,#1(! ,%! -/L#! ,-#! 0/9#! #$$#2,! /0! ,-#!
/9%3(,! %$! :&%,#1(! :&%)32#)! .4! ,5%! 2%:1#0! %$! ,-/,! /**#*#6! ! 7%&! ,-#!
9/_%&1,4! %$! <#(#0! 0,3)1#)+! ,-#! (%&9/*! Y16#6! .*"'",<=#Z! /**#*#0! /&#!
-/:*%03$$121#(,+!0%!#L#(!1$!/!93,/,1%(!2/30#0!/!2%9:*#,#!*%00!%$!$3(2,1%(!
1(! %(#! /**#*#+! ,-#! 51*)=,4:#! /**#*#! 51**! .#! )%91(/(,! /()! ,-#! (%&9/*!
$3(2,1%(! %$! ,-#! .1%2-#912/*! :/,-5/4! 51**! .#! &#,/1(#)6! ! ! Q(! ,-#! %,-#&!
-/()+! 1(! 0%9#! .1%2-#912/*! :/,-5/40+! /! 01(<*#! 51*)=,4:#! /**#*#! 9/4! .#!
0!="1*&$522*>*#&,+!16#6!1,!9/4!(%,!:&%)32#!#(%3<-!:&%,#1(!,%!&#03*,!1(!/!
(%&9/*!:-#(%,4:#!1$!,-#!%,-#&!/**#*#!1(!/!-#,#&%K4<%,#!10!(%,!$3(2,1%(/*6!!
G(! ,-10! 2/0#+! /! (%(=$3(2,1%(/*! /**#*#! 51**! .#! )%91(/(,! ,%! /! 51*)=,4:#!
/**#*#6!

IWA!XRBBAII!F`R@HA!@BC!'QBQWV"HGC!NHQFFAF!
P#(#,1210,0+!1(2*3)1(<!'#()#*+!9/;#!30#!%$!,+5#!D+##'*&7!"*&#$!Y71<3&#!
O6a!/Z6!!I-#0#!/&#!:%:3*/,1%(0!%$!:*/(,0!%&!/(19/*0!1(!5-12-!/**!:/&#(,0!
/()! ,-#1&! %$$0:&1(<! -/L#! 1)#(,12/*! :-#(%,4:#0! %L#&! 9/(4! <#(#&/,1%(0!
51,-! &#0:#2,! ,%! /! :/&,123*/&! ,&/1,6! I&3#! .&##)1(<! *1(#0! /&#! 30#$3*+!
.#2/30#! ,-#4! 2/(! .#! /0039#)! ,%! .#! -%9%K4<%30! $%&! ,-#! /**#*#0! ,-/,!
/$$#2,!/!,&/1,!%$!1(,#&#0,6!J-#(!,5%!1()1L1)3/*0!,-/,!/&#!-%9%K4<%30!$%&!
,-#! 0/9#! /**#*#0! /&#! 2&%00#)+! /**! %$! ,-#1&! %$$0:&1(<! 51**! /**! /*0%! .#!
-%9%K4<%306!

!
Q ( # ! D % 2 3 0 !S!8"E!!

!
!
Figure 3.6 a) a true- P1L#(! ,-#! <#(%,4:#0! %$! /(4! ,5%! :/&#(,0+! 5#! 2/(! :&#)12,! /**! %$! ,-#!
breeding line b) a :%001.*#!<#(%,4:#0!%$!,-#!%$$0:&1(<6!!73&,-#&9%&#+!1$!5#!/*0%!;(%5!,-#!
monohybrid cross )%91(/(2#! &#*/,1%(0-1:0! $%&! /**! %$! ,-#! /**#*#0+! 5#! 2/(! :&#)12,! ,-#!
produced by mating
two pure-breeding
:-#(%,4:#0! %$! ,-#! %$$0:&1(<6! ! @! 2%(L#(1#(,! 9#,-%)! $%&! 2/*23*/,1(<! ,-#!
lines #8:#2,#)! <#(%,4:12! /()! :-#(%,4:12! &/,1%0! $&%9! /! 2&%00! 10! ,-#! 30#! %$!
X3((#,,!FT3/&#+!5-12-!10!/!9/,&18!1(!5-12-!/**!%$!,-#!:%001.*#!</9#,#0!
:&%)32#)!.4!%(#!:/&#(,!/&#!*10,#)!/*%(<!%(#!/810+!/()!,-#!</9#,#0!$&%9!
,-#! %,-#&! :/&#(,! /&#! *10,#)! /*%(<! ,-#! %,-#&! /8106! ! A/2-! :%001.*#!
2%9.1(/,1%(! %$! </9#,#0! 10! *10,#)! /,! ,-#! 1(,#&0#2,1%(! %$! #/2-! &%5! /()!
2%*39(6!!!

J-#(! ,5%! -#,#&%K4<%,#0! /&#! 2&%00#)+! 5-/,! <#(%,4:#0! /&#! :&%)32#)+!

/()! 5-/,! 10! ,-#! #8:#2,#)! $&#T3#(24! %$! #/2-! <#(%,4:#?! ! ! G$! 5#! 30#! ,-#!
049.%*0!!!/()!"!,%!&#:&#0#(,!#/2-!,-#!,5%!/**#*#0!%$!,-#!-#,#&%K4<%,#+!
,-#(!$&%9!,-#!F5&&#,,!645!+#!Y71<3&#!O6bZ!5#!0##!,-/,!,-&##!<#(%,4:#0!
/&#! :&%)32#)! 1(! /! &/,1%! %$! [McM[6! ! G$! 5#! ;(%5! 0%9#,-1(<! /.%3,! ,-#!
:-#(%,4:#0!/()!)%91(/(2#!&#*/,1%(0-1:0!%$!,-#0#!/**#*#0!Y/0!19:*1#)!.4!
,-#!049.%*0!30#)Z+!5#!2/(!:&#)12,!,-#!#8:#2,#)!:-#(%,4:12!&/,1%!%$!,-#!
:&%<#(4! $&%9! ,-10! 2&%00! 10! OM[6! ! @! 2&%00! .#,5##(! ,5%! 1()1L1)3/*0! ,-/,!
/&#! .%,-! -#,#&%K4<%30! /,! /! 01(<*#! *%230! 10! 0%! 2%99%(! 1(! <#(#,120! ,-/,!
,-10!2&%00!10!<1L#(!1,0!%5(!(/9#M!!,-#!:1&10<D+*'!>+1$$!Y71<3&#!O6a!.Z6!!!

! !! "

!! !! !"
Figure 3.7 A Punnett
Square showing a
"! !" ""
monohybrid cross
!

!
 Q ( # ! D % 2 3 0 !S!8"G!!

IAFI!NHQFFAF!N@B!"A!RFAC!IQ!CAIAH'GBA!PABQIVXAF!
d(%51(<!,-#!<#(%,4:#0!%$!/(!1()1L1)3/*!10!303/**4!/(!19:%&,/(,!:/&,!%$!
/!<#(#,12!#8:#&19#(,6!!W%5#L#&+!<#(%,4:#0!2/((%,!.#!%.0#&L#)!)1&#2,*4>!
,-#4!930,!.#!1($#&&#)!./0#)!%(!:-#(%,4:#06!"#2/30#!%$!)%91(/(2#+!1,!10!
%$,#(! (%,! :%001.*#! ,%! )10,1(<310-! .#,5##(! /! -#,#&%K4<%,#! /()! /!
-%9%K<4%,#!./0#)!%(!:-#(%,4:#!/*%(#!Y#6<6!0##!,-#!:3&:*#=$*%5#&#)!7c!
:*/(,0!1(!71<3&#!O6a.Z6!!I%!)#,#&91(#!,-#!<#(%,4:#+!/!,#$,!>+1$$!2/(!.#!
:#&$%&9#)+! 1(! 5-12-! /(! 1()1L1)3/*! 51,-! /(! /9.1<3%30! <#(%,4:#! 10!
2&%00#)! 51,-! /(! 1()1L1)3/*! ,-/,! 10! -%9%K4<%30! &#2#001L#! $%&! /**! %$! ,-#!
*%21! .#1(<! ,#0,#)6! ! 7%&! #8/9:*#+! 1$! 4%3! 5#&#! <1L#(! /! :#/! :*/(,! 51,-!
:3&:*#! $*%5#&0! Y/()! (%! %,-#&! 1($%&9/,1%(Z+! 4%3! 5%3*)! (%,! ;(%5!
5-#,-#&!,-#!:*/(,!5/0!/!-%9%K4<%,#!Y!!Z!%&!-#,#&%K4<%,#!Y!"Z6!!!V%3!
2%3*)! 2&%00! ,-10! :3&:*#=$*%5#&#)! :*/(,! ,%! /! 5-1,#=$*%5#&#)! :*/(,! /0! /!
,#$,#++! 01(2#! 4%3! ;(%5! ,-#! <#(%,4:#! %$! ,-#! ,#0,#&! 10! ""6! ! V%3! 51**!
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! ! !

!!"! !" !" Figure 3.8 A Punnett

Squares showing
!!"! !" !" examples of test
crosses.
!

! ! "

"! !" ""

"! !" ""

FAU=DGBd@PA!GF!@B!G'XQHI@BI!AUNAXIGQB!IQ!'ABCADEF!7GHFI!
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U!/()!V6!!

Figure 3.7 Reciprocal

crosses involving a
sex-linked trait.

!
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,-#(!0#:/&/,#*4!2&%00!/!&#)=#4#)!$#9/*#!51,-!/!5-1,#=#4#)!9/*#6!!B%,#!
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*1(;#)! <#(#0! /&#! )1$$#&#(,! $&%9! 5-/,! 10! #8:#2,#)! $%&! /3,%0%9/*! <#(#0!
Y71<3&#!O6eZ6!!

!
! #$%! #$% #$ #$
Figure 3.8 Reciprocal !
crosses involving a #!$! +$#+!$ +$#+!$ #!$% +$+!$# +$+$#!
sex-linked trait. !
&!! +$#,! +$#, &! +$, +$,!
!

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Figure 3.9 Probability
distribution of the chi-
square statistic for five
different degrees of
freedom

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A1A1 A1A2 A2A2

1 all hairs black on the same all hairs white
individual:
50% of hairs are all
black
50% of hairs are all
white
2 all hairs black all hairs are the same all hairs white
shade of grey
3 all hairs black all hairs black 50% of individuals
have all white hairs

50% of individuals
have all black hairs

4 all hairs black all hairs black mice have no hair

5 all hairs black all hairs white all hairs white
6 all hairs black all hairs black all hairs white
7 all hairs black all hairs black hairs are a wide range
of shades of grey
!

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 C h a p t e r   4  | 4­5  

Box 4‐1 Transposable Elements 
Transposable  elements  (TEs)  occur  naturally  throughout  the  chromosomes  of  almost  all 
organisms.  These DNA sequences have a unique ability to be inserted into new locations in the 
genome,  after  being  cut  or  copied  from  their  original  location.      TEs  are  also  known  as  mobile 
genetic  elements,  or  more  informally  as  jumping  genes.    The  locations  into  which  TEs  are 
inserted are not entirely random, but TEs can in principle be inserted into almost any region of 
the  genome.    TEs  can  therefore  move  into  other  genes,  causing  mutations.    Researchers  have 
methods of artificially increasing the rate of transposition, making TEs a useful type of mutagen.  
However the biological importance of TEs extends far beyond their use in mutant screening; TEs 
are  also  important  causes  of  disease  and  phenotypic  instability,  and  they  are  a  major  force  in 
evolution. 

There  are  two  major  classes  of  TEs  in  eukaryotes  (Figure  4‐B1).    Class  I  elements  include 
retroposons  and  retrotransposons;  these  are  copied  by  means  of  an  RNA  intermediate.  The 
transcript  is  reverse  transcribed  into  DNA  before  being  inserted  elsewhere  in  the  genome 
through  the  action  of  enzymes  such  as  integrase.    Class  II  elements  are  known  also  as 
transposons; these do not use reverse transcriptase or an RNA intermediate for transposition.  
Using an enzyme called transposase, most transposons are cut from their original location and 
then this excised dsDNA fragment is inserted into a new location. Note that the name transposon 
is sometimes used incorrectly to refer to any type of TEs, but in this book we use transposon to 
refer only to Class II elements.  

 
Figure  4­B1  .    Representative  examples  of  the  two  main  types  of  transposable  elements.  (TEs)  
Class  I  elements  transpose  via  an  ssRNA  intermediate,  which  is  reverse  transcribed  to  dsDNA 
prior  to  insertion  of  this  copy  in  a  new  site  in  the  genome.  Class  II  elements  do  not  involve  an 
RNA intermediate; most Class II elements are cut from their original location as dsDNA, prior to 
being inserted into a new site in the genome.  Although the diagram shows TEs being inserted on 
the same chromosome as they originated from, TEs can also move to other chromosomes within 
the same cell. 

  
C h a p t e r   4  | 4­6  

TEs are relatively short DNA sequences (between 100bp and 10kb), and encode no more than a 
few proteins (if any).   Normally, the protein‐coding genes within a TE are all related to the TE’s 
own  transposition  functions,  e.g.  reverse  transcriptase,  transposase,  and  integrase.  However, 
some TEs (of either Class I or II) do not encode any proteins at all.  These non­autonomous TEs 
can  only  transpose  if  they  are  supplied  with  enzymes  produced  by  other,  autonomous  TEs 
located  elsewhere  in  the  genome.      In  all  cases,  enzymes  for  transposition  recognize  conserved 
nucleotide sequences within the TE, which show the enzymes where to begin cutting or copying, 
and re‐insertion.   

The human genome is nearly 45% TEs, the vast majority of which are families of Class I elements 
called LINEs and SINEs.  The short, Alu type of SINE occurs in more than one million copies in 
the human genome (compare this to the approximately 30,000, non‐TE, protein‐coding genes in 
humans).      Indeed,  TEs  make  up  a  significant  portion  of  the  genomes  of  almost  all  eukaryotes.  
Class  I  elements,  which  usually  transpose  via  a  copy‐and‐paste  mechanism,  tend  to  be  more 
abundant  than  Class  II  elements,  which  mostly  use  a  cut‐and‐paste  mechanism.      But  even  the 
cut‐paste  mechanism  can  lead  to  an  increase  in  TE  copy  number,  in  some  circumstances  (for 
example,  if  the  site  vacated  by  the  excised  transposon  is  repaired  with  a  DNA  template  from  a 
homologous chromosome that itself contains a copy of a transposon).  

Besides greatly expanding the DNA content of genomes, TEs contribute to genome evolution in 
many  other  ways.    As  already  mentioned,  they  may  disrupt  gene  function  by  insertion  into  a 
gene’s coding region or regulatory region.  More interestingly adjacent regions of chromosomal 
DNA are sometimes mistakenly transposed along with the TE; this can lead to gene duplication.  
The  duplicated  genes  are  then  free  to  evolve  independently,  leading  in  some  cases  to  the 
development  of  new  functions.    The  breakage  of  strands  by  TE  excision  and  integration  can 
disrupt genes, and can lead to chromosome rearrangement or deletion if errors are made during 
strand  rejoining.  Furthermore,  having  so  many  similar  TE  sequences  distributed  throughout  a 
chromosome  sometimes  allows  mispairing  of  regions  of  homologous  chromosomes  at  meiosis, 
which can cause unequal crossing‐over, resulting in deletion or duplication of large segments of 
chromosomes.   Thus, TEs are an important evolutionary force, and are not merely “junk DNA” as 
they were once called.  

   

Figure  4­B2.    Barbara  McClintock  won  a  Nobel  Prize  for  her  discovery  of  TEs.    She  did  so  by 
studying pigment variegation in maize kernels, which is caused by the movement of TEs in and 
out of pigmentation genes.  This is an example of phenotypic instability. 
  

 
 C h a p t e r   4  | 4­7  

The new strand therefore bears a mutation, which will be inherited, in 
all the strands that are subsequently replicated from it. 

Figure 4.6 Alkylation of

guanine (shown in red)
allows G to bond with
thymine rather than cytosine
at replication.

 
Intercalating  agents  are  another  type  of 
chemical  mutagen.    These  induce  mutations  by 
inserting  between  the  stacked  bases  at  the 
center  of  the  DNA  helix  (Figure  4.7).    This 
intercalation distorts the shape of the DNA helix, 
which can cause the wrong bases to be added to 
a  growing  DNA  strand  during  DNA  synthesis.  
Intercalating  agents  tend  to  be  flat,  planar 
molecules  such  as  benzopyrene,  a  component 
of wood and tobacco smoke.  Another important 
intercalating  agent  is  thalidomide,  an  anti‐
nausea  drug  whose  harmful  effects  were 
unknown until its consumption by thousands of 
Figure 4.7 Benzopyrene
pregnant  woman  resulted  in  birth  defects.   (circled in red) is an
Finally  ethidium  bromide,  the  dye  that  example of an
fluorescently stains DNA in laboratory assays, is  intercalating agent
also  an  intercalating  agent.    For  this  reason, 
molecular biologists are trained to handle this chemical carefully. 

PHYSICAL 
Anything  that  damages  DNA  by  transferring  energy  to  it  can  be 
considered  a  physical  mutagen.    Usually  this  involves  radioactive 
particles,  x‐rays,  or  UV  light.    Smaller,  fast  moving  particles  may 
substitute or delete a single base, while larger, slightly slower particles 
induce larger deletions by breaking the double stranded helix.  Physical 

  
C h a p t e r   4  | 4­8  

mutagens  can  also  create  unusual  structures  in  DNA,  such  as  the 
thymine  dimers  formed  by  UV  light  (Figure  4.8).  Thymine  dimers 
disrupt  normal  base‐pairing  in  the  double  helix,  and  may  block 
replication altogether if not repaired by the cell. 

Figure 4.8 Thymine

dimers are formed
when adjacent thymine
bases on the same
DNA strand become
covalently linked (red
bonds) follow exposure
to mutagens such as
UV light. The dimers
distort base pairing and
can interrupt processes
such as replication.

MUTANT SCREENING: FORWARD GENETICS 
One way to identify genes that affect a particular biological process is to 
induce random mutations in a population, and then look for individuals 
with  phenotypes  that  might  be  caused  by  a  disruption  of  a  particular 
biochemical  pathway.    This  strategy  of  mutant  screening  has  been 
used very effectively to better understand the molecular components of 
hundreds  of  different  biological  processes.        For  example,  to  better 
understand  processes  of  memory  and  learning,  researchers  have 
screened  mutagenized  populations  of  Drosophila  to  identify  flies  (or 
larvae)  that  lack  the  normal  ability  to  learn  to  associate  a  particular 
odor with an electric shock.  Some of the genes identified by this mutant 
screen  may  be  relevant  to  learning  and  memory  in  other  animals, 
including conditions such as Alzheimer’s disease in humans.  

Exposure of an organism to a mutagen causes mutations in essentially 
random  positions  along  the  chromosomes.    Most  of  the  mutant 
phenotypes  recovered  from  a  genetic  screen  are  caused  by  loss­of­
function mutations.  These are changes in the sequence of an allele that 
cause  it  to  no  longer  produce  the  same  level  of  active  protein  as  the 
wild‐type  allele.    Loss‐of‐function  alleles  tend  to  be  recessive  because 
the  wild‐type  allele  is  haplosufficient  (see  Chapter  6).        A  loss‐of‐
function allele that produces no active protein is called an amorph, or 
null.  On the other hand, alleles with only a partial loss‐of‐function are 
called hypomorphic.     More rarely, a mutant allele may have a gain­of­
function,  producing  either  more  of  the  active  protein  (hypermorph) 
or producing an active protein with a new function (neomorph).  

  

 
 C h a p t e r   4  | 4­9  

In a typical mutant screen, researchers will treat a parental population 
with  a  mutagen.  This  may,  for  example,  involve  soaking  seeds  in  EMS, 
or mixing this mutagen with the food fed to flies.  The individuals that 
are directly exposed with the mutagen are called the M0 generation.  No 
phenotypes will be visible among the M0 generation, in part because not 
all  somatic  (i.e.  non‐reproductive)  cells  of  the  M1  individuals  will  be 
affected in the same way.  More important in the M1 generation are the 
germline  cells:  gametes  and  any  of  their  developmental  precursors. 
Since a single gamete contributes half of the chromosomes to every cell 
of  the  next  (M2)  generation,  mutagenizing  gametes  or  the  cells  that 
produce them is the easiest way to generate individuals in which every 
cell  bears  the  same  mutation.      In  most  cases,  however,  the  M1 
individual  will  be  heterozygous  for  any  induced  mutation;  this  is 
because  it  would  be  very  unlikely  for  the  same  gene  to  have  been 
mutated  in  the  other  gamete  that  contributed  to  the  M1  individual.  
Because most induced mutations are recessive, the M1  generation must 
therefore  be  selfed  (i.e.  crossed  to  siblings,  or  self‐fertilization  if  the 
species  is  a  hermaphrodite  like  worms  and  most  plants),  and  the 
following generations (e.g. M2,  in which mutant alleles could potentially 
become homozygous) examined for the presence of novel phenotypes. 
Once  a  relevant  mutant  has  been  identified,  geneticists  can  begin  to 
make inferences about what the normal function of the mutated gene is, 
based on its mutant phenotype.   

SOME MUTATIONS MAY NOT HAVE DETECTABLE PHENOTYPES 
The vast majority of mutations (especially substitutions) have no effect 
on  the  phenotype.    Often,  this  is  because  the  mutation  is  silent;  it 
changes  the  DNA  sequence  of  a  non‐coding  region  of  the  DNA,  or  else 
the changes a base within a codon without changing the amino acid that 
it encodes (recall that the genetic code is degenerate; for example, GCT, 
GCC,  GCA,  and  GCG  all  encode  alanine).    There  are  also  cases  where  a 
mutation  can  cause  a  complete  loss‐of‐function  of  a  gene,  yet  not 
produce a phenotype even when the mutant allele is homozygous.  This 
can  often  be  attributed  to  genetic  redundancy,  i.e.  the  encoding  of 
similar  genes  at  more  than  one  locus  in  the  genome.    It  is  important 
therefore to remember this important limitation of mutational analysis:  
genes  with  redundant  functions  cannot  be  easily  identified  by  mutant 
screening. 

Some  phenotypes  require  individuals  to  reach  a  particular 

developmental  stage  before  they  can  be  scored.    For  example,  flower 
color  can  only  be  scored  in  plants  that  are  mature  enough  to  make 
flowers,  and  eye  color  can  only  be  scored  in  flies  that  have  developed 
eyes.  However, some alleles may not develop sufficiently to be included 
among  the  progeny  that  are  scored  according  to  their  phenotype.   
Lethal  alleles  that  arrest  the  development  of  an  individual  at  an 
embryonic stage may therefore go unnoticed in a typical mutant screen.  
Furthermore,  the  progeny  of  a  monohybrid  cross  involving  an 
embryonic  lethal  recessive  allele  may  therefore  all  be  of  a  single 

  
C h a p t e r   4  | 4­10  

phenotypic class, giving a phenotypic ratio of 1:0 (which is the same as 
3:0). 

Many  genes  are  first  identified  in  mutant  screens,  and  so  they  tend  to 
be  named  after  their  mutant  phenotypes.    This  can  cause  some 
confusion  for  students  of  genetics.    For  example,  we  have  already 
encountered an X‐linked gene named white in fruit flies.  Null mutants 
of white have white eyes, but the normal function of the white gene is 
actually the production of a red pigment.   

COMPLEMENTATION TESTING 
 

Mutations  in  different  genes  can  produce  the  same  phenotype.    For 
example, in the biochemical pathway shown in Figure 4.9, a plant that 
lacks  the  function  of  gene  A  (genotype  aa)  would  produce  mutant, 
white flowers that looked just like the flowers of a plant that lacked the 
function of gene B (genotype bb).  The genetics of two loci are discussed 
more in the following chapters. 

Figure 4.9 In this

simplified biochemical
pathway, two enzymes
encoded by two
different genes modify
chemical compounds in
two sequential
reactions to produce a
purple pigment. Loss
of either of the
enzymes disrupts the
pathway and no
pigment is produced.

 
 

As explained earlier in this chapter, mutant screening is one of the main 
activities of geneticists.  When geneticists find two mutants with similar 
phenotypes, either in natural populations or during a mutant screen, an 
immediate  question  is  whether  or  not  the  mutants  have  lost  the 
function of the same gene.  The other possibility is that each mutant has 
  

 
 C h a p t e r   4  | 4­11  

lost  the  function  of  a  different  gene.    If  they  are  both  mutants  of  the 
same gene, then their genotypes can both be represented as aa.  If each 
has a mutation in a different gene, then the genotype of one individual 
can  be  represented  as  aa,  and  the  other  individual  as  bb  (or  more 
completely as aaBB and AAbb, respectively).   

A technique called complementation testing can be used to determine 
whether  two  individuals  with  the  same  phenotype  carry  mutations  in 
the same gene or different genes.  The only requirement of this test is 
two  pure‐breeding  individuals  with  the  same  phenotype;  no  prior 
knowledge  of  the  genes  or  biochemical  pathways  is  required.      To 
perform  a  complementation  test,  two  homozygous  individuals  with 
similar mutant phenotypes are crossed (Figure 4.10).   If the F1 progeny 
all have the mutant phenotype, then we infer that same gene is mutated 
in each parents.  If the F1 progeny all appear to be wild‐type, then each 
of the parents most likely carries a mutation in a different gene (Figure 
4.11). 

Figure 4.10
In a typical
complementation test,
the genotypes of two
parents are unknown
(although they must be
pure breeding,
homozygous mutants).
If the F1 progeny all
have a mutant
phenotype there (Case
1), there is no
complementation. If
the F1 progeny are all
wild-type, the
mutations are said to
have complemented
each other. See Figure
4.11 for interpretation.  
 

  
C h a p t e r   4  | 4­12  

Figure 4.11  
The pure breeding,
homozygous mutants
parents had unknown
genotypes before the
complementation test, but
it could be assumed that
they were either mutations
in the same genes (Case 1)
or different genes (Case
2). In Case 1, all of the
progeny would have a
mutant phenotype,
because they would all
have the same,
homozygous genotype as
the parents. In Case 2,
each parent has a mutation
in a different gene,
therefore none of the F1
progeny will be  
homozygous mutant at
any one locus. Note that  
the genotype in Case 1
could be written as either  
aa or aaBB.
 

Thus, if two homozygous mutants produce F1 progeny that have a wild‐
type  phenotype,  complementation  is  said  to  have  occurred,  because 
the wild‐type alleles of each of the genes were able to compensate for 
the  recessive,  mutant  alleles  that  were  also  inherited  (Case  2,  Figure 
4.11).      On  the  other  hand,  if  the  F1  progeny  all  have  a  mutant 
phenotype,  (case  1,  Figure  4.11),  the  mutant  genotypes  fail  to 
complement  each  other,  because  they  contain  mutations  of  the  same 
gene. These could be either the same mutant alleles, or different mutant 
alleles of the same gene. If the two mutants are independent (e.g. they 
came  from  different  natural  populations  or  from  independently 
mutagenized  individuals),  the  mutations  are  probably  different  alleles 
of the same gene.   All mutants that fail to complement each other are 
said to be in the same complementation group.  

  

 
 C h a p t e r   4  | 4­13  

_______________________________________________________________________________________________________________ 

SUMMARY 
• When a variation in DNA sequence originated recently, and is rare in a population, we call 
that change a mutation. 
 
• When  variations  in  DNA  sequence  co‐exist  in  a  population,  and  neither  one  can  be 
meaningfully defined as wild‐type, we call the variations polymorphisms. 
 
• Mutations may either occur spontaneously, or may be induced by exposure to mutagens. 
 
 
• Mutations may result in either substitutions, deletions, or insertions. 
 
• Mutation  usually  causes  either  a  partial  or  complete  loss  of  function,  but  sometimes 
results in a gain of function,  including new functions. 
 
• Spontaneous  mutations  arise  from  many  sources  including  natural  errors  in  DNA 
replication, usually associated with base mispairing, or else insertion deletion especially 
within repetitive sequences. 
 
• Induced  mutations  result  from  mispairing,  DNA  damage,  or  sequence  interruptions 
caused by chemical, biological, or physical mutagens.  
 
• By randomly inducing mutations, then screening for a specific phenotype, it is possible to 
identify genes associated with specific biological pathways.  
 
• Transposable  elements  are  dynamic,  abundant  components  of  eukaryotic  genomes  and 
important forces in evolution. 
 
• Transposable  elements  are  dynamic,  abundant  components  of  eukaryotic  genomes  and 
important forces in evolution. 
 
• Mutation of different genes can produce a similar phenotype. 
 
• Complementation testing determines whether two mutants are the result of mutation of 
the same gene, or if each mutant is caused by mutation of a different gene. 
 
• The  efficiency  of  mutant  screening  is  limited  by  silent  mutations,  redundancy,  and 
embyronic lethality. 

  
C h a p t e r   4  | 4­14  

KEY TERMS 
mutation  transposon   
mutant  retrotransposon  loss‐of‐function 
polymorphism  reverse transcriptase  gain‐of‐function 
insertion  transposase  amorph 
deletion  non‐autonomous  null 
substitution  autonomous  hypomorph 
mutagen  SINE, LINE, Alu  hypermorph 
biological mutagen  P‐element  neopmorph 
chemical mutagen  T‐DNA  somatic 
physical mutagen  alkylation agent  germline 
tautomer  EMS  M0, M1, M2 
mispairing  intercalating agent  silent mutation 
loop  benzopyrene  redundancy 
SSR  ethidium bromide  lethality 
insertional mutagen  thymine dimer  complementation group 
Class I, Class II  mutant screen   
 
_______________________________________________________________________________________________________________

 
STUDY QUESTIONS 
4.1  How  are  polymorphisms  and  mutations  However there are twice as many females as 
alike?  How are they different?  males in the F1 progeny of this cross. 
   a)  Propose  a  hypothesis  to  explain  these 
4.2  What  are  all  of  the  ways  a  substitution  observations. 
can occur in a DNA sequence?  b) How could you test your hypothesis? 
   
4.3  What  are  all  of  the  ways  a  deletion  can  4.7 You decide to use genetics to investigate 
occur in a DNA sequence?  how your favourite plant makes its 
  flowers smell good. 
4.4 What are all of the ways an insertion can  a) What steps will you take to identify some 
occur in a DNA sequence?  genes that are required for production of the 
  sweet floral scent? Assume that this plant 
4.5 In the context of this chapter, explain the  is a self‐pollinating diploid. 
health hazards of smoking tobacco.  b)  One  of  the  recessive  mutants  you 
  identified has fishy‐smelling flowers, so you 
4.6 You have exposed a female fruit fly to a  name  the  mutant  (and  the  mutated  gene) 
mutagen.  Mating  this  fly  with  a  non‐ fishy.    What  do  you  hypothesize  about  the 
mutagenized  male  produces  offspring  that  normal function of the wild‐type fishy gene? 
appear to be completely normal. 

  

 
 

c)  Another  recessive  mutant  lacks  floral   

scent altogether, so you call it nosmell. What  4.10  You  are  interested  in  finding  genes 
could you hypothesize about the normal  involved  in  synthesis  of  proline  (Pro),  an 
function of this gene?  amino acid that is normally synthesizes by a 
  particular model organism.   
4.8  Suppose  you  are  only  interested  in  a)  How would you design a  mutant screen 
finding dominant mutations that affect floral  to    identify  genes  required  for  Pro 
scent.  synthesis?  
a)  What  do  you  expect  to  be  the  relative  b)  Imagine  that  your  screen  identified  ten 
frequency  of  dominant  mutations,  as  mutants  (#1  through  #4)  that  grew  poorly 
compared to recessive mutations, and why?  unless  supplemented  with  Pro.    How  could 
b)  How  will  you  design  your  screen  you  determine  the  number  of  different 
differently  than  in  the  previous  question,  in  genes represented by these mutants? 
order  to  detect  dominant  mutations  c)  If  each of the four mutants represents a 
specifically?  different gene, what will be the phenotype of 
c)  Which  kind  of  mutagen  is  most  likely  to  the F1 progeny if  any pair of the four 
produce  dominant  mutations,  a  mutagen  mutants are crossed? 
that produces point mutations, or a mutagen  d)  If  each of the four mutants represents 
that produces large deletions?  the same gene, what will be the phenotype 
  of the F1 progeny if  any pair of the four 
  mutants are crossed?
4.9  Which  types  of  transposable  elements 
are transcribed?  
 

 
M#&,-$1!N >?@ABC??H!DE@!>O>PFD"AOEH!

Figure 5.1 Polydactyly

is an example of a
human trait that can be
studied by pedigree
analysis
!!
"#$!%&'()!)*+)$,-'!*.!/$+$-()'!0$')1(%$0!(+!-#$!,1$)$0(+/!)#&,-$1'!)&+!
%$! &,,2($0! -*! &23*'-! &+4! $56&14*-()! *1/&+('37! ! 8*9$:$1;! '*3$!
-$)#+(<5$';! '5)#! &'! -$'-! )1*''$';! )&+! *+24! %$! ,$1.*13$0! 9(-#! 3*0$2!
*1/&+('3'!*1!*-#$1!',$)($'!-#&-!)&+!%$!$=,$1(3$+-&224!3&+(,52&-$07!"*!
'-504! -#$! (+#$1(-&+)$! ,&--$1+'! *.! /$+$'! (+! #53&+'! &+0! *-#$1! ',$)($'!!
.*1! 9#()#! )*+-1*22$0! 3&-(+/'! &1$! +*-! ,*''(%2$;! /$+$-()('-'! 5'$!
-$)#+(<5$'!(+)250(+/!-#$!&+&24'('!*.!,$0(/1$$'!&+0!,*,52&-(*+'!

>?@ABC??!DEDFGHAH!
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(+0(:(05&2J'! '$=;! .&3(24! 1$2&-(*+'#(,'! &+0! ,#$+*-4,$7! "#$'$! 0(&/1&3'!
&1$!5'$0!-*!0$-$13(+$!-#$!!"#$!"%!&'($)&*+',$!*.!&!,&1-()52&1!0('$&'$!
*1! -1&(-;! &+0! -*! ,1$0()-! -#$! ,1*%&%(2(-4! *.! (-'! &,,$&1&+)$! &3*+/!
*..',1(+/7!!>$0(/1$$!&+&24'('!('!-#$1$.*1$!&+!(3,*1-&+-!-**2!(+!%*-#!%&'()!
1$'$&1)#!&+0!-$'$*&,!,".'/$0&'-7!

?&)#! ,$0(/1$$! 1$,1$'$+-'! &22! *.! -#$! &:&(2&%2$! (+.*13&-(*+! &%*5-! -#$!
(+#$1(-&+)$!*.!&!'(+/2$!-1&(-!K3*'-!*.-$+!&!0('$&'$L!9(-#(+!&!.&3(247!!"#$!
,$0(/1$$! ('! -#$1$.*1$! 01&9+! .1*3! .&)-5&2! (+.*13&-(*+! 1&-#$1! -#&+!
-#$*1$-()&2!,1$0()-(*+';!%5-!-#$1$!('!&29&4'!'*3$!,*''(%(2(-4!*.!$11*1'!(+!
 > $ 0 ( / 1 $ $ ' ! Q ! > * , 5 2 & - ( * + ' ! R!1"2!!

-#('! (+.*13&-(*+;! $',$)(&224! 9#$+! 1$24(+/! *+! .&3(24! 3$3%$1'J!

Figure 5.2 Symbols
used in drawing a
pedigree.
1$)*22$)-(*+'! *1! $:$+! )2(+()&2! 0(&/+*'$'7! ! A+! 1$&2! ,$0(/1$$';! .51-#$1!
)*3,2()&-(*+'!)&+!&1('$!05$!-*!(+)*3,2$-$!,$+$-1&+)$!K(+)250(+/!&/$!*.!
*+'$-L!&+0!:&1(&%2$!$=,1$''(:(-4!*.!0('$&'$!&22$2$';!%5-!.*1!-#$!$=&3,2$'!
,1$'$+-$0! (+! -#('! %**6;! 9$! 9(22! ,1$'53$! )*3,2$-$! &))51&)4! *.! -#$!
,$0(/1$$'7! ! D! ,$0(/1$$! 3&4! %$! 01&9+! 9#$+! -14(+/! -*! 0$-$13(+$! -#$!
+&-51$! *.! &! +$924! 0(')*:$1$0! 0('$&'$;! *1! 9#$+! &+! (+0(:(05&2! 9(-#! &!
.&3(24!#('-*14!*.!&!0('$&'$!9&+-'!-*!6+*9!-#$!,1*%&%(2(-4!*.!,&''(+/!-#$!
0('$&'$!*+!-*!-#$(1!)#(201$+7!!A+!$(-#$1!)&'$;!&!-1$$!('!01&9+;!&'!'#*9+!(+!
S(/51$!N7T;!9(-#!)(1)2$'!-*!1$,1$'$+-!.$3&2$';!&+0!'<5&1$'!-*!1$,1$'$+-!
3&2$'7!!A.!&+!(+0(:(05&2!('!6+*9+!-*!#&:$!'43,-*3'!*.!-#$!0('$&'$;!-#$!
'43%*2!('!.(22$0!(+7!!H*3$-(3$'!&!#&2.U.(22$0!(+!'43%*2!('!5'$0!-*!(+0()&-$!
&! ,+))&$)! *.! &! 0('$&'$V! -#('! ('! '*3$*+$! 9#*! 0*$'! +*-! #&:$! &+4!
'43,-*3'!*.!-#$!0('$&'$;!%5-!9#*!,&''$0!-#$!0('$&'$!*+!-*!'5%'$<5$+-!
/$+$1&-(*+'7! ! E*-$! -#&-! 9#$+! &! ,$0(/1$$! ('! )*+'-15)-$0;! (-! ('! *.-$+!
5+6+*9+!9#$-#$1! &!,&1-()52&1!(+0(:(05&2!('!&!)&11($1!*1!+*-;!'*!+*-! &22!
)&11($1'!&1$!&29&4'!$=,2()(-24!(+0()&-$0!(+!&!,$0(/1$$7!!S*1!'(3,2()(-4;!9$!
9(22! &''53$! -#&-! -#$! ,$0(/1$$'! ,1$'$+-$0! (+! -#('! )#&,-$1! &1$! &))51&-$;!
&+0!1$,1$'$+-!.5224!,$+$-1&+-!-1&(-'7!!
!

AES?CCAEB!"8?!WO@?!OS!AE8?CA"DEM?!
B(:$+!&!,$0(/1$$!*.!&+!5+)#&1&)-$1(I$0!0('$&'$!*1!-1&(-;!*+$!*.!-#$!.(1'-!
-&'6'! ('! -*! 0$-$13(+$! -#$! 3*0$! *.! (+#$1(-&+)$;! &'! -#('! (+.*13&-(*+! ('!
$''$+-(&2!(+!)&2)52&-(+/!-#$!,1*%&%(2(-4!-#&-!-#$!-1&(-!9(22!%$!(+#$1(-$0!(+!
*..',1(+/7!!X$!9(22!)*+'(0$1!.*51!3&Y*1!-4,$'!*.!(+#$1(-&+)$Z!&5-*'*3&2!
0*3(+&+-! KD@L;! &5-*'*3&2! 1$)$''(:$! KDCL;! [U2(+6$0! 0*3(+&+-! K[@L;! [U
2(+6$0!1$)$''(:$!K[CL7!!!

!"#$%$&!'!($&)*!*#!+!(,!
!

Figure 5.3 A pedigree

consistent with AD
inheritance.
!
X#$+!&!0('$&'$!('!)&5'$0!%4!&!0*3(+&+-!&22$2$!*.!&!/$+$;!$:$14!,$1'*+!
9(-#!-#&-!&22$2$!9(22!'#*9!'43,-*3'!*.!-#$!0('$&'$!K&''53(+/!)*3,2$-$!
,$+$-1&+)$L;! &+0! *+24! *+$! 0('$&'$! &22$2$! +$$0'! -*! %$! (+#$1(-$0! .*1! &+!
(+0(:(05&2!-*!%$!&..$)-$07!!!"#5';!$:$14!&..$)-$0!(+0(:(05&2!35'-!#&:$!&+!
&..$)-$0! ,&1$+-7! ! D! ,$0(/1$$! 9(-#! &..$)-$0! (+0(:(05&2'! (+! $:$14!
/$+$1&-(*+! ('! -4,()&2! *.! D@! 0('$&'$'7! ! 8*9$:$1;! %$9&1$! -#&-! *-#$1!
3*0$'!*.!(+#$1(-&+)$!)&+!&2'*!'#*9!-#$!0('$&'$!(+!$:$14!/$+$1&-(*+;!&'!
0$')1(%$0! %$2*97! ! A-! ('! &2'*! ,*''(%2$! .*1! &+! &..$)-$0! (+0(:(05&2! 9(-#! &+!
!

!
 > $ 0 ( / 1 $ $ ' ! Q ! > * , 5 2 & - ( * + ' !R!1"3!!

D@!0('$&'$!-*!#&:$!&!.&3(24!9(-#*5-!&+4!&..$)-$0!)#(201$+;!(.!-#$!&..$)-$0!
,&1$+-! ('! &! #$-$1*I4/*-$7! ! ! "#('! ('! ,&1-()52&124! -15$! (+! '3&22! .&3(2($';!
9#$1$!-#$!,1*%&%(2(-4!*.!$:$14!)#(20!(+#$1(-(+/!-#$!+*13&2;!1&-#$1!-#&+!
0('$&'$!&22$2$!('!+*-!$=-1$3$24!'3&227!!E*-$!-#&-!D@!0('$&'$'!&1$!5'5&224!
1&1$!9(-#(+!,*,52&-(*+';!-#$1$.*1$!&..$)-$0!(+0(:(05&2'!9(-#!D@!0('$&'$'!
-$+0! -*! %$! #$-$1*I4/*-$'! K*-#$19('$;! %*-#! ,&1$+-'! 9*520! #&:$! #&0! -*!
%$$+!&..$)-$0!9(-#!-#$!'&3$!1&1$!0('$&'$L7!!D)#*+01*,2&'-()!09&1.('3;!
&+0! ,*240&)-424! &1$! %*-#! $=&3,2$'! *.! #53&+! )*+0(-(*+'! -#&-! 3&4!
.*22*9!&+!D@!3*0$!*.!(+#$1(-&+)$7!

-"')*./(!($&)*!*#!+-(,! Figure 5.4 Two

! pedigrees consistent with
XD inheritance.

!
!

A+![U2(+6$0!0*3(+&+-!(+#$1(-&+)$;!-#$!/$+$!1$',*+'(%2$!.*1!-#$!0('$&'$!
('!2*)&-$0!*+!-#$![U)#1*3*'*3$;!&+0!-#$!&22$2$!-#&-!)&5'$'!-#$!0('$&'$!
('!0*3(+&+-!-*!-#$!+*13&2!&22$2$7!!\$)&5'$!.$3&2$'!#&:$!-9()$!&'!3&+4!
[U)#1*3*'*3$'!&'!3&2$';!.$3&2$'!-$+0!-*!%$!&..$)-$0!3*1$!.1$<5$+-24!
-#&+! 3&2$'! (+! [@! ,$0(/1$$'7! ! 8*9$:$1;! +*-! &22! ,$0(/1$$'! ,1*:(0$!
'5..()($+-! (+.*13&-(*+! -*! 0('-(+/5('#! [@! &+0! D@7! O+$! 0$.(+(-(:$!
(+0()&-(*+! -#&-! &! -1&(-! ('! (+#$1(-$0! &'! D@! 1&-#$1! -#&+! [@! ('! -#&-! &+!
&..$)-$0!.&-#$1!,&''$'!-#$!0('$&'$!-*!&!'*+V!-#('!-4,$!*.!-1&+'3(''(*+!('!
+*-! ,*''(%2$! 9(-#! [@;! '(+)$! 3&2$'! (+#$1(-! -#$(1! [! )#1*3*'*3$! .1*3!
-#$(1!3*-#$1'7!

!"#$%$&!'!0/1/%%)2/!+!0,!
@('$&'$'! -#&-! &1$! (+#$1(-$0! (+! &+! &5-*'*3&2! 1$)$''(:$! ,&--$1+! 1$<5(1$!
-#&-!%*-#!,&1$+-'!*.!&+!&..$)-$0!(+0(:(05&2!)&114!&-!2$&'-!*+$!)*,4!*.!-#$!!
0('$&'$! &22$2$7! ! ! X(-#! DC! -1&(-';! 3&+4! (+0(:(05&2'! (+! &! ,$0(/1$$! )&+! %$!
)&11($1';! ,1*%&%24! 9(-#*5-! 6+*9(+/! (-7! ! M*3,&1$0! -*! ,$0(/1$$'! *.!
0*3(+&+-!-1&(-';!DC!,$0(/1$$'!-$+0!-*!'#*9!.$9$1!&..$)-$0!(+0(:(05&2'!
&+0! &1$! 3*1$! 2(6$24! -#&+! D@! *1! [@! -*! ]'6(,! &! /$+$1&-(*+^7! ! "#5';! -#$! Figure 5.5 Some types
3&Y*1! .$&-51$! -#&-! 0('-(+/5('#$'! DC! .1*3! D@! *1! [@! ('! -#&-! 5+&..$)-$0! of rickets may follow an
(+0(:(05&2'!)&+!#&:$!&..$)-$0!*..',1(+/7! XD mode of inheritance.
 > $ 0 ( / 1 $ $ ' ! Q ! > * , 5 2 & - ( * + ' ! R!1"4!!

Figure 5.6 A pedigree

consistent with AR
inheritance.
!
!
Figure 5.7 Many inborn
errors of metabolism,
such as phenylketonuria
(PKU) are inherited as
AR. Newborns are often
tested for a few of the
most common metabolic
diseases.

-"')*./(!0/1/%%)2/!+-0,!
\$)&5'$!3&2$'!#&:$!*+24!*+$![U)#1*3*'*3$;!&+4!3&2$!-#&-!(+#$1(-'!&+!
[U2(+6$0! 1$)$''(:$! 0('$&'$! &22$2$! 9(22! %$! &..$)-$0! %4! (-! K&''53(+/!
)*3,2$-$! ,$+$-1&+)$L7! ! ! "#$1$.*1$;! (+! [C! 3*0$'! *.! (+#$1(-&+)$;! 3&2$'!
-$+0!-*!%$!&..$)-$0!3*1$!.1$<5$+-24!-#&+!.$3&2$'7!!"#('!('!(+!)*+-1&'-!-*!
DC! &+0! D@;! 9#$1$! %*-#! '$=$'! -$+0! -*! %$! &..$)-$0! $<5&224;! &+0! [@;! (+!
9#()#!.$3&2$'!&1$!&..$)-$0!3*1$!.1$<5$+-247!!E*-$;!#*9$:$1;!-#&-!(+!-#$!
'3&22!'&3,2$!'(I$'!-4,()&2!*.!#53&+!.&3(2($';!(-!3&4!+*-!%$!,*''(%2$!-*!
&))51&-$24!0$-$13(+$!9#$-#$1!*+$!'$=!('!&..$)-$0!3*1$!.1$<5$+-24!-#&+!
*-#$1'7!!!O+!-#$!*-#$1!#&+0;!*+$!.$&-51$!*.!&!,$0(/1$$!-#&-!)&+!%$!5'$0!-*!
0$.(+(-(:$24! $'-&%2('#! -#&-! &+! (+#$1(-&+)$! ,&--$1+! ('! +*-! [C! ('! -#$!
,1$'$+)$!*.!&+!&..$)-$0!0&5/#-$1!.1*3!5+&..$)-$0!,&1$+-'V!%$)&5'$!'#$!
9*520!#&:$!#&0!-*!(+#$1(-!*+$![U)#1*3*'*3$!.1*3!#$1!.&-#$1;!#$!9*520!
&2'*!#&:$!%$$+!&..$)-$0!(+![C7!!

!
 > $ 0 ( / 1 $ $ ' ! Q ! > * , 5 2 & - ( * + ' !R!1"1!!

Figure 5.8 A pedigree consistent with XR inheritance. Figure 5.9 Some forms of colour
blindness are inherited as XR-traits.
!! Colour blindness is diagnosed using tests
such as this Ishihara Test.
H>OCD@AM!DE@!EOEU8?CA"D\F?!@AH?DH?H!!
O+24!&!.1&)-(*+!*.!&22!*.!-#$!6+*9+!#53&+!-1&(-'!&+0!0('$&'$'!#&:$!%$$+!
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#&:$! &! #$1(-&%2$! )*3,*+$+-;! %5-! #&:$! 3*1$! )*3,2$=! (+#$1(-&+)$!
,&--$1+'! 05$! -*! -#$! (+:*2:$3$+-! *.! 352-(,2$! /$+$'! &+0! $+:(1*+3$+-&2!
.&)-*1'7!!!O+!-#$!*-#$1!#&+0;!'*3$!0('$&'$'!3&4!&,,$&1!-*!%$!#$1(-&%2$;!
%$)&5'$!-#$4!&..$)-!352-(,2$!3$3%$1'!*.!-#$!'&3$!.&3(24;!%5-!-#('!)*520!
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*-#$1! $+:(1*+3$+-&2! .&)-*1'! (+! -#$(1! #*3$'7! ! S(+&224;! 0('$&'$'! 9(-#!
'(3(2&1! '43,-*3'! 3&4! #&:$! 0(..$1$+-! )&5'$';! '*! *.! 9#()#! 3&4! %$!
/$+$-()!9#(2$!*-#$1'!&1$!+*-7!!O+$!$=&3,2$!*.!-#('!('!DFH!K&34*-1*,#()!
2&-$1&2!')2$1*'('LV!&,,1*=(3&-$24!NU_`a!*.!)&'$'!&1$!(+#$1(-$0!(+!&+!D@!
,&--$1+;! 9#(2$! -#$! 3&Y*1(-4! *.! -#$! 1$3&(+(+/! )&'$'! &,,$&1! -*! %$!
/5")+#&,;! (+! *-#$1! 9*10';! +*-! )&5'$0! %4! &! 35-&-(*+! (+#$1(-$0! .1*3! &!
,&1$+-7!!!X$!+*9!6+*9!-#&-!0(..$1$+-!/$+$'!*1!,1*-$(+'!&1$!&..$)-$0!(+!
-#$! (+#$1(-$0! &+0! ',*1&0()! .*13'! *.! DFH7! "#$! ,#4'()('-! H-$,#$+! Figure 5.10
8&96(+/! &+0! %&'$%&22! ,2&4$1! F*5! B$#1(/! %*-#! '5..$1$0! .1*3! ',*1&0()! Stephen Hawking
DFH7!!

MDFMPFD"AEB!>CO\D\AFA"A?H!
O+)$! -#$! 3*0$! *.! (+#$1(-&+)$! *.! &! 0('$&'$! *1! -1&(-! ('! 6+*9+;! '*3$!
(+.$1$+)$'!&%*5-!-#$!/$+*-4,$!*.!(+0(:(05&2'!(+!&!,$0(/1$$!)&+!%$!3&0$;!
%&'$0! *+! -#$(1! ,#$+*-4,$'7! ! B(:$+! -#$'$! /$+*-4,$';! (-! ('! ,*''(%2$! -*!
)&2)52&-$! -#$! ,1*%&%(2(-4! *.! &! ,&1-()52&1! /$+*-4,$! %$(+/! (+#$1(-$0! (+!
'5%'$<5$+-! /$+$1&-(*+'7! ! "#('! )&+! %$! 5'$.52! (+! /$+$-()! )*5+'$2(+/;! .*1!
$=&3,2$!9#$+!,1*',$)-(:$!,&1$+-'!9('#!-*!6+*9!-#$!2(6$2(#**0!*.!-#$(1!
*..',1(+/!(+#$1(-(+/!&!0('$&'$!.*1!9#()#!-#$4!#&:$!&!.&3(24!#('-*147!

>1*%&%(2(-($'!(+!,$0(/1$$'!&1$!)&2)52&-$0!5'(+/!-#$!'&3$!%&'()!3$-#*0'!
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