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4.1
Protozoa
Metazoa
4.2
Fertilized egg
Two-cell stage(cleavage begins)
Four-cell stage
Eight-cell stage
Radial Cleavage Spiral Cleavage
4.3
Cnidaria: Aurelia
Annelida: Eupomatus
Cephalochordata: Brachiostoma
4.4
1 2 3 4 5 6 7 8 9 10 11 12 13
Anterior Medial Posterior
a-1 a-13
b-1 b-9
c-4 c-13
d-1 d-13
Vertebrates
1 10
Branchiostoma floridae(Chordata)
lab pb Dfd SerAntpUbx AbdA
AbdB
Drosophila melanogaster(Arthropoda)
Drosophila pseudoobscura(Arthropoda)
mxp Dfd Cx Ptl Utx A Extra urogomphiTribolium castaneum(Arthropoda)
AfDfdAfSer
AfAntpAfUbx
AfabdAArtemia(Arthropoda)
1 8Ctenodrilus serratus(Annelida)
lin-39(ceg-15)
egl-5(ceg-11)
ceh-13 mab-5Caenorhabditis elegans(Nemertina)
lab ?pb Dfd AntpSchistosoma mansoni(Platyhelminthes)
Cnox1 Cnox2 antpC
Chlorohydra viridissima(Cnidaria)
4.5
Fruit fly Ubx(Insect)
Mouse Ubx(Mammal)
C. elegans Ubx(Nematoda)
Leech Ubx(Annelid)
Amphioxus pb(Urochordate)
Frog AbdB(Amphibian)
old
ems
lab pb Dfd Scr Ubx AbdB eve
The Zootype
4.6
Strain: Ork
0
20
40
60
80
100
120
140
0 2 4 6 8
Added Salt (%)
Per
cent
age
Sur
viva
l A
rea
of A
nal P
apilla
e
Strain: sp2bs2
0
20
40
60
80
100
120
140
0 2 4 6 8
Added Salt (%)
Strain: al b c sp2
0
20
40
60
80
100
120
140
0 2 4 6 8
Added Salt (%)
4.7
2.00
1.75
1.50
1.25
1.00
0.75
197
3
198
3
199
3
Year 200
3 0.2 0.1 0.0
-0.1 -0.2
Plasticity
Fit
nes
s
4.8
Original phenotypes
Phenotypicchanges
Changes inStructural genes
New environments
are entered
Natural selection acts on new
phenotypic traits
Changes in diet, daily patterns of activity, social interactions, etc.
Reproductive isolation
4.9
HETEROCHRONY
Different morphologyDifferent body size
Same morphologyDifferent body size
Peramorphosis
D develops new stages not present
in A.
Paedomorphosis
D adult similar to juvenile A.
Predisplacement
Onset of developmentoccurs earlier in D,
relative to A.
Acceleration
Juvenile developmentis accelerated in D,
relative to A.
Peramorphosis
D develops new stages not present
in A.
Postdisplacement
Onset of developmentis delayed in D,
relative to A.
Neoteny
Juvenile development is retarded in D,
relative to A.
Progenesis
Sexual maturity occurs earlier in D than in A.
Gigantism
D larger than A in body size.
Dwarfism
D smaller than A in body size.
4.10
(a) (b)
4.11
A B C
Fertilization Sexual maturity
B C D
Predisplacement
A B C
A B C
A
Acceleration
Hypermorphosis
A B C D
A B C
Fertilization Sexual maturity
A B
Postdisplacement
A B C
Neoteny
A B
A B C
Progenesis
A B
(b) Peramorphosis(a) Paedomorphosis
Descendant
Descendant
Descendant
Ancestor
Ancestor
Ancestor
B C D
4.12
Ho
rn L
eng
th (
mm
)
Basilar Length of Skull (mm)
600
400
200
100
60
40
20
10 8
6
4
400 1000
4.13
• Early response to external stimuli
• Delayed sensitive period for socialization
Behavior
• Early sexual maturation
• Longer breeding season
• Larger litter size
Physiology
• Floppy ears
• Shorter or rolled tail
• Short legs
• Depigmentation
Morphology
Artificial selection target
HormonesNeurotransmitters
Tamability
Developmental program (simplified)
Genes
Correlated changes
4.14
Att
ack
Fre
qu
ency
S1
S4
S13
20
30
10
0
20
30
10
0
20
30
10
0
40 80 160 180 220 260
Age (Days)
4.15
(a) (b)
(c) (d)
(e)
4.16
Telencephalon
Procencephalon Diencephalon
Mesencephalon
Rhombencephalon
A4 A3 A2 A1
Hox A Genes
4.17
(a) (b)
4.18
Pairs of littermates
Weaned at about 21-24 days of age
Randomly assigned
Enriched environment
Impoverished environment
Treatment duration(e.g., 30 days)
Behavioraltesting
Chemical and morphological
analyses of the CNS