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Tom Wenseleers University of Leuven, Belgium Ph.D. defence May 22nd, 2001
Conflict from Cell to Colony
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Cooperation is Key Feature in
Evolution of Life on Earth
Genes to Genomes
Prokaryotes to Eukaryotes
Unicellular to Multicellular Organisms
Organisms to Societies
Major transitions in evolution
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Cooperation seems obvious to explain when viewed in terms of species-level benefits
But erroneous logic: non-cooperative ’free-riders’ outcompete altruists
But potential for conflict
Potential for Conflict in
Most Societies
Conflicts may occur between organisms, but also between cells or genes (’intragenomic conflict’)
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In what ratio should males and females
be reared?F
½
M½ ¼
M¾
F
Conflicts in insect societies
Equal Sex-Ratio
Equal Sex-Ratio
3:1 FemaleBiased
Sex-Ratio
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Cytoplasmic sex-ratio distorters
Conflict also occurs at the genomic level: maternally transmitted genes favour more female biased sex-ratios than nuclear genes(“intragenomic conflict”)
Cytoplasmic genes such as mitochondria or some bacterial symbionts may manipulate host to produce female biased broods (“cytoplasmic sex-ratio distorters”)
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Wolbachia Example of a maternally
transmitted symbiont
Alpha-proteobacterium
Occurs mainly in arthropods (insects+Crustacea) + nematodes
Manipulates host reproduction to favour own spread
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FemaleBiased
Sex-Ratios
Male Killing
Feminisation
Parthenogenesis Induction
Cytoplasmic Incompatibility
Effects on host reproduction
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NormalOffspring
Production
Reduces fitness of Uninfected Female x Infected Male Crosses
Gives an advantage to infected females
Sterility in diploids, but production
of males only in haplo-diploids
Cytoplasmic incompatibility
Inviable
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Phylogeny
Oth
er a
lpha
pro
teobacte
ria
Ehrlichieae
Neorickettsia
Gamm
a
prote
obac
teria
0.1
Wolbachia
CaedibacterMtK
MitochondriaCMS
Orientia MK
Rickettsia MK
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Aims of my thesis Part I : empirical
– Does Wolbachia occur in ant societies?
– Alternative explanation for female biased sex-ratios in this group?
Part II : theoretical– What do animal and genomic
conflicts have in common?– Can sociobiological theory be
applied to both?
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S e q u e n c e o f E v e n t s
Modelling
Make predictions
DNA Analysis
Measure key parameters
Experiments
Formally test hypotheses
Ideas Hypotheses
Molecular Data
Experimental Data
Integrated approach
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Part I. Wolbachia - a cause of intragenomic conflict
in ant colonies
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Work plan
Does Wolbachia occur in ant societies and if so in what frequency?
What effects does it have?Three case studies :– Parthenogenetic species– Wood ant Formica truncorum– Leptothorax nylanderi
Host-parasite coevolution?
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Polymerase Chain Reaction using Specific Primers
Targets: ftsZ and wsp Wolbachia genes
Positive, negative and nuclear DNA (18S rDNA) controls
Negative samples retested twice
Methodology: PCR Assay
Sensitive& Reliable
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High Incidence Worldwide
Indonesia
Chapter 1Wenseleers et al. (1998) Proceedings of the Royal Society of London
# species=50
Florida
Jeyaprakash & Hoy (2000) Insect Molecular Biology
# species=10
Panama
Van Borm et al. (2001) Journal of Evolutionary Biology
# species=7
Europe
# species=50 Chapter 6
3451 samples
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Morphological evidence
Present in trophocytes and oocytes
Electron and light microscopical (DAPI) evidence
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Work plan
Does Wolbachia occur in ant societies and if so in what frequency? YES, IN HIGH FREQUENCY
What effects does it have?Three case studies :– Parthenogenetic species– Wood ant Formica truncorum– Leptothorax nylanderi
Host-parasite coevolution?
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Work plan
Does Wolbachia occur in ant societies and if so in what frequency? YES, IN HIGH FREQUENCY
What effects does it have?Three case studies :– Parthenogenetic species– Wood ant Formica truncorum– Leptothorax nylanderi
Host-parasite coevolution?
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Parthenogenesis induction?
Were not infected.Parthenogenesis not induced by
Wolbachia.
PCR Assay
Grasso et al. (2000) Ethology, Ecology & Evolution 12:309-314Wenseleers & Billen (2000) Journal of Evolutionary Biology 13:277-280
6 Parthenogenetic Antsand Cape Honey Bee
N=25036 cols.
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Wolbachia in F. truncorum
With: Lotta Sundström University of Helsinki
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Formica truncorum
Extensive variation in sex-ratio produced by different colonies
Linked to facultative sex-ratio biasing :– Workers kill brothers in colonies
headed by singly mated queen– But not in colonies with double
mated queen
Does Wolbachia affect the sex-ratio too?
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Effect on the sex-ratio :
– Males should be infected less than queens
– Sex-ratio should be correlated with infection rates
Incompatibility :
– Males and queens should be infected equally
– Uninfected colonies should not be able to survive
Predictions
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Formica truncorum
Males (96%) and queens (94%) infected equally
All colonies infected (total # 33) despite production of 6% uninfected queens by each colony
Consistent with an incompatibility effect :
Uninfected queens do not survive past the founding stage due to incompatible matings
Wenseleers, Sundström & Billen (2002) Proceedings of the Royal Society of London series B, in press.
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r2 = 0.0097
0
0.25
0.5
0.75
1
0.00 0.20 0.40 0.60 0.80 1.00
Percent infected workers
Inve
stm
ent i
n fe
mal
es
GLM Effects F p
No. of mates 4.88 0.04Infection rate 0.85 0.37Colony size 0.69 0.42
Infection and sex-ratio
Wenseleers, Sundström & Billen (2002) Proceedings of the Royal Society of London series B, in press.
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r2 = 0.03
0
4
8
12
0.00 0.20 0.40 0.60 0.80 1.00
Proportion infected adult workers
Per
cap
ita
pro
du
ctio
n
Worker production
r2 = 0.28
0
4
8
12
0.00 0.20 0.40 0.60 0.80 1.00
Proportion infected adult workers
Per
cap
ita
pro
du
ctio
n
Sexual production
GLM
Effects F p F p
No. of mates 2.11 0.16 2.5 0.13
Infection rate 2.89 0.11 10.2 0.005
Infection and colony fitness
Wenseleers, Sundström & Billen (2002) Proceedings of the Royal Society of London series B, in press.
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p<0.015p<0.0001
0
25
50
75
100
Pe
rce
nt
infe
cte
d
Sexuals Adult workers
Worker pupae
Infection rates
N=296 N=158 N=387
Adaptiveclearance to
reduce colony load?
Wenseleers, Sundström & Billen (2002) Proceedings of the Royal Society of London series B, in press.
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Conclusions
No effects on the sex-ratio
Probably causes incompatible matings
Deleterious effects on colony function, but partly mitigated by clearance of infection in adult workers
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Leptothorax nylanderi
Test experimentally whether Wolbachia causes incompatible matings
Setup: antibiotic treatment as an artificial means of creating the uninfected queen x infected male crossing type
Prediction: male production (infertility) following antibiotic treatment
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0.4
0.5
0.6
0.7
0.8
0.9
1
Untreated Treated
Pri
ma
ry s
ex
-ra
tio
2 = 10.51, p < 0.001
Antibiotics experiments
4 coloniesN=70
7 coloniesN=152
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Work plan
Does Wolbachia occur in ant societies and if so in what frequency? YES, IN HIGH FREQUENCY
What effects does it have?Three case studies :– Parthenogenetic species– Wood ant Formica truncorum– Leptothorax nylanderi
Host-parasite coevolution?
![Page 31: Tom Wenseleers University of Leuven, Belgium Ph.D. defence May 22nd, 2001 Conflict from Cell to Colony](https://reader031.vdocument.in/reader031/viewer/2022012913/56649e895503460f94b8dc3b/html5/thumbnails/31.jpg)
Wolbachia surface protein wsp was sequenced (approx. 550 bp)
Direct cycle sequencing when ants were infected by single strain
Cloning and sequencing when ants were infected by multiple strains (TA-cloning kit, pUC57 vector)
Methodology: Sequencing
28 sequencesAligned with previously sequenced relatives
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So
len
op
sis
invi
cta
(im
po
rted
) C
ole
om
egill
a m
acu
lata
len
gi
Dia
ph
ori
na
citr
i P
lute
lla x
ylo
stel
la L
aod
elp
hax
str
iate
llus
Acr
aea
ence
do
n 1
Tri
cho
pri
a T
sp2
Dry
inid
was
p sp
Por
celli
onid
es p
ruin
osus
Sph
aero
ma
rugi
caud
a
Bac
toce
ra c
ucur
bita
e
Trib
oliu
m m
aden
s
Trib
olium
confu
sum
Rhin
ophoridae
unid
Doro
nomyrmex kutte
ri B
Doro
nomyrmex pacis B
2
Trichogramma spp.
Adalia bipunctata B
Coleomegilla maculata
Adalia bipunctata A
Acromyrmex octospinosus B3
Acromyrmex insinuator B1
Acromyrmex echinatior B
Solenopsis invicta (native)
Acromyrmex octospinosus B1 Acromyrmex octospinosus B2 Acromyrmex insinuator B2
Myrmica sabuleti Telenomus nawai Encarsia formosa
Diplolepis rosae
Leptopilina australis
Cadra cautella
Tetranychus urticae
Acraea encedon
Culex quinquefasciatus
Culex pipiens (ESPRO)
Drosophila simulans (W
atsonville)
Aedes albopictus (Houston)
Doronom
yrmex pacis B
1
Isopods
Trichopria drosophilae
Asobara tabida
Myrm
ica sulcin
od
is (Sam
so D
)
Myrm
ica sulcin
od
is (Ru
ssia)
Teleu
tom
yrmex sch
neid
eri
Neo
chryso
charis fo
rmo
sa
Fo
rmic
a ru
faD
acu
s d
esti
llato
ria
Do
ron
om
yrm
ex g
oes
swal
di A
2
Do
ron
om
yrm
ex p
acis
A4
Do
ron
om
yrm
ex k
utt
eri A
Fo
rmic
a fu
sca
(Mo
ls D
)
Fo
rmic
a fu
sca
(SJW
B)
Fo
rmic
a fu
sca
(KH
B)
Lep
toth
ora
x ac
ervo
rum
Bac
toce
ra s
p 1
Asc
D
Cat
agly
phis
iber
ica
Glo
ssin
a au
sten
i
Form
ica
poly
cten
a
Form
ica
trunc
orum
Formic
a pra
tensi
s
Asobara t
abida 3
Drosophila
sechellia
Drosophila sim
ulans (Hawaii)
Cadra cautella 2
Doronomyrmex pacis A3
Gnamptogenys menadensis
Phlebotomus papatasi (Israel)
Doronomyrmex goesswaldi A1
Acromyrmex octospinosus A1Solenopsis invicta A (native)Doronomyrmex pacis A2
Solenopsis richteri A
Acromyrmex echinatior A1
Drosophila simulans (Riverside)
Drosophila melanogaster (CantonS)
Drosophila melanogaster (Cairns)
Drosophila simulans (Coffs Harbour)
Aedes albopictus (Houston)
Nasonia vitripennis A
Drosophila bifasciata
Glossina morsitans centralis
Leptopilina heterotoma 2
Trichogramm
a bourarachae
Trichogramm
a kaykai (LC110)
Muscidifurax uniraptor
Acrom
yrmex insinuator A
Plagiolepis pygmaea
Myrm
ica sulcinodis (Pyrenees)
Formica lem
ani
Myrm
ica rub
ra
Do
ron
om
yrmex p
acis A1
0.050(25 MY)
A B
High strain diversity
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So
len
op
sis
invi
cta
(im
po
rted
) C
ole
om
egill
a m
acu
lata
len
gi
Dia
ph
ori
na
citr
i P
lute
lla x
ylo
stel
la L
aod
elp
hax
str
iate
llus
Acr
aea
ence
do
n 1
Tri
cho
pri
a T
sp2
Dry
inid
was
p sp
Por
celli
onid
es p
ruin
osus
Sph
aero
ma
rugi
caud
a
Bac
toce
ra c
ucur
bita
e
Trib
oliu
m m
aden
s
Trib
olium
confu
sum
Rhin
ophoridae
unid
Doro
nomyrmex kutte
ri B
Doro
nomyrmex pacis B
2
Trichogramma spp.
Adalia bipunctata B
Coleomegilla maculata
Adalia bipunctata A
Acromyrmex octospinosus B3
Acromyrmex insinuator B1
Acromyrmex echinatior B
Solenopsis invicta (native)
Acromyrmex octospinosus B1 Acromyrmex octospinosus B2 Acromyrmex insinuator B2
Myrmica sabuleti Telenomus nawai Encarsia formosa
Diplolepis rosae
Leptopilina australis
Cadra cautella
Tetranychus urticae
Acraea encedon
Culex quinquefasciatus
Culex pipiens (ESPRO)
Drosophila simulans (W
atsonville)
Aedes albopictus (Houston)
Doronom
yrmex pacis B
1
Isopods
Trichopria drosophilae
Asobara tabida
Myrm
ica sulcin
od
is (Sam
so D
)
Myrm
ica sulcin
od
is (Ru
ssia)
Teleu
tom
yrmex sch
neid
eri
Neo
chryso
charis fo
rmo
sa
Fo
rmic
a ru
faD
acu
s d
esti
llato
ria
Do
ron
om
yrm
ex g
oes
swal
di A
2
Do
ron
om
yrm
ex p
acis
A4
Do
ron
om
yrm
ex k
utt
eri A
Fo
rmic
a fu
sca
(Mo
ls D
)
Fo
rmic
a fu
sca
(SJW
B)
Fo
rmic
a fu
sca
(KH
B)
Lep
toth
ora
x ac
ervo
rum
Bac
toce
ra s
p 1
Asc
D
Cat
agly
phis
iber
ica
Glo
ssin
a au
sten
i
Form
ica
poly
cten
a
Form
ica
trunc
orum
Formic
a pra
tensi
s
Asobara t
abida 3
Drosophila
sechellia
Drosophila sim
ulans (Hawaii)
Cadra cautella 2
Doronomyrmex pacis A3
Gnamptogenys menadensis
Phlebotomus papatasi (Israel)
Doronomyrmex goesswaldi A1
Acromyrmex octospinosus A1Solenopsis invicta A (native)Doronomyrmex pacis A2
Solenopsis richteri A
Acromyrmex echinatior A1
Drosophila simulans (Riverside)
Drosophila melanogaster (CantonS)
Drosophila melanogaster (Cairns)
Drosophila simulans (Coffs Harbour)
Aedes albopictus (Houston)
Nasonia vitripennis A
Drosophila bifasciata
Glossina morsitans centralis
Leptopilina heterotoma 2
Trichogramm
a bourarachae
Trichogramm
a kaykai (LC110)
Muscidifurax uniraptor
Acrom
yrmex insinuator A
Plagiolepis pygmaea
Myrm
ica sulcinodis (Pyrenees)
Formica lem
ani
Myrm
ica rub
ra
Do
ron
om
yrmex p
acis A1
0.050(25 MY)
A B
No match with host phylogeny
Acrom
yrmex insinuator A
Plagiolepis pygmaea
Myrm
ica sulcinodis (Pyrenees)
Formica lem
ani
Myrm
ica rub
ra
Do
ron
om
yrmex p
acis A1
Hosts diverged 35 MY ago, but share a recently evolved W. strain
(1.7 MY old)
Doro
nomyrmex kutte
ri B
Doro
nomyrmex pacis B
2
Doronom
yrmex pacis B
1
Do
ron
om
yrm
ex g
oes
swal
di A
2
Do
ron
om
yrm
ex p
acis
A4
Do
ron
om
yrm
ex k
utt
eri ADoronomyrm
ex pacis A3Doronomyrmex goesswaldi A1
Doronomyrmex pacis A2
Do
ron
om
yrmex p
acis A1
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So
len
op
sis
invi
cta
(im
po
rted
) C
ole
om
egill
a m
acu
lata
len
gi
Dia
ph
ori
na
citr
i P
lute
lla x
ylo
stel
la L
aod
elp
hax
str
iate
llus
Acr
aea
ence
do
n 1
Tri
cho
pri
a T
sp2
Dry
inid
was
p sp
Por
celli
onid
es p
ruin
osus
Sph
aero
ma
rugi
caud
a
Bac
toce
ra c
ucur
bita
e
Trib
oliu
m m
aden
s
Trib
olium
confu
sum
Rhin
ophoridae
unid
Doro
nomyrmex kutte
ri B
Doro
nomyrmex pacis B
2
Trichogramma spp.
Adalia bipunctata B
Coleomegilla maculata
Adalia bipunctata A
Acromyrmex octospinosus B3
Acromyrmex insinuator B1
Acromyrmex echinatior B
Solenopsis invicta (native)
Acromyrmex octospinosus B1 Acromyrmex octospinosus B2 Acromyrmex insinuator B2
Myrmica sabuleti Telenomus nawai Encarsia formosa
Diplolepis rosae
Leptopilina australis
Cadra cautella
Tetranychus urticae
Acraea encedon
Culex quinquefasciatus
Culex pipiens (ESPRO)
Drosophila simulans (W
atsonville)
Aedes albopictus (Houston)
Doronom
yrmex pacis B
1
Isopods
Trichopria drosophilae
Asobara tabida
Myrm
ica sulcin
od
is (Sam
so D
)
Myrm
ica sulcin
od
is (Ru
ssia)
Teleu
tom
yrmex sch
neid
eri
Neo
chryso
charis fo
rmo
sa
Fo
rmic
a ru
faD
acu
s d
esti
llato
ria
Do
ron
om
yrm
ex g
oes
swal
di A
2
Do
ron
om
yrm
ex p
acis
A4
Do
ron
om
yrm
ex k
utt
eri A
Fo
rmic
a fu
sca
(Mo
ls D
)
Fo
rmic
a fu
sca
(SJW
B)
Fo
rmic
a fu
sca
(KH
B)
Lep
toth
ora
x ac
ervo
rum
Bac
toce
ra s
p 1
Asc
D
Cat
agly
phis
iber
ica
Glo
ssin
a au
sten
i
Form
ica
poly
cten
a
Form
ica
trunc
orum
Formic
a pra
tensi
s
Asobara t
abida 3
Drosophila
sechellia
Drosophila sim
ulans (Hawaii)
Cadra cautella 2
Doronomyrmex pacis A3
Gnamptogenys menadensis
Phlebotomus papatasi (Israel)
Doronomyrmex goesswaldi A1
Acromyrmex octospinosus A1Solenopsis invicta A (native)Doronomyrmex pacis A2
Solenopsis richteri A
Acromyrmex echinatior A1
Drosophila simulans (Riverside)
Drosophila melanogaster (CantonS)
Drosophila melanogaster (Cairns)
Drosophila simulans (Coffs Harbour)
Aedes albopictus (Houston)
Nasonia vitripennis A
Drosophila bifasciata
Glossina morsitans centralis
Leptopilina heterotoma 2
Trichogramm
a bourarachae
Trichogramm
a kaykai (LC110)
Muscidifurax uniraptor
Acrom
yrmex insinuator A
Plagiolepis pygmaea
Myrm
ica sulcinodis (Pyrenees)
Formica lem
ani
Myrm
ica rub
ra
Do
ron
om
yrmex p
acis A1
0.050(25 MY)
A B
Multiple infections
Doro
nomyrmex pacis B
2
Doronom
yrmex pacis B
1Do
ron
om
yrm
ex p
acis
A4
Doronomyrmex pacis A3
Doronomyrmex pacis A2
Do
ron
om
yrmex p
acis A1
Multi infections may drive speciation
events!
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No match with host phylogeny
pratensis
lemani
fusca
rufa
O
100
99
polyctena
truncorum84100
0.02(10 MY)
...and their symbionts
rufa
polyctena
pratensis
truncorum
lemani
fusca
O
Formica hosts...
Gyllenstrand, unpublished
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Work plan
Does Wolbachia occur in ant societies and if so in what frequency? YES, IN HIGH FREQUENCY
What effects does it have?Three case studies :– Parthenogenetic species– Wood ant Formica truncorum– Leptothorax nylanderi
Host-parasite coevolution? NO, OCCASIONAL HORIZONTAL TRANSMISSION
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Part II. Theoretical aspects ofconflict and cooperation
With: Francis Ratnieks and Kevin Foster
University of Sheffield
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Animal vs. intragenomic conflict
What do animal and intragenomic conflict have in common?
Is there a “general theory of conflict” that provides insight into the evolution of conflict at both levels?
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Theories of conflict
Two Approaches in the Study of Conflict
Kin SelectionHamilton
Game Theoryvon Neumann &
Morgenstern
Single method
r.B > CCost Dependson Social Context
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Generalised Hamilton’s rule
Wenseleers & Ratnieks submitted
j jgB.r - C+E . > 0β
Hamilton’s rule(costs & benefits
independent of social context)
Terms thattake into
account socialcontext
Consequence ofboth cooperating
Regression of genotype on joint behaviour
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Animal vs. intragenomic conflict
0 -B
B -C
DOVE HAWK
DO
VE
HA
WKANIMAL CONFLICT
COOPERATEDRIVE
CO
OP
ER
ATE
DR
IVE
GDC.(1-k)1/2
GDC.k GDD/2
GENOMIC CONFLICT(MEIOTIC DRIVE)
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Animal vs. intragenomic conflict
Shows that game theoretic logic of conflict at both levels is the same
But can genes also be related?
Yes, kinship measures genetic correlation and for 2 genes at a locus this is the inbreeding coefficient FIT
When genes are related they are selected to be altruistic !
Application of generalised Hamilton’s rule allows detailed analysis
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Spite: Hamilton’s unproven theory
Medea killed her children to take away the smile from her husband’s face.
Example of a paradoxical behaviour that harms another at no benefit to self (“spite”)
We showed that some forms of intragenomic conflict qualify as spiteful behaviour (Maternal effect lethals, queen killing in the fire ant)
Foster, Ratnieks & Wenseleers (2000) Trends in Ecology & Evolution 15:469-470Foster, Wenseleers & Ratnieks (2001) Annales Zoologici Fennici, in press
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Why become a worker? Why do social insect females work
for the benefit of others?
Usual explanation: indirect genetic benefit when altruism is directed towards relatives (’kin selection’)
But is relatedness in insect societies high enough?
E.g. honey bee: queen mates with several males so that workers mostly rear half-sisters (r=0.3)
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New calculations Female should become a queen with a
probability of (1-Rf)/(1+Rm) (self determination)
– = 20% for stingless bees (singly mated)
– = 56% for honey bees (polyandrous)
Too high for the colony as a whole, since queens are only needed for swarming (“tragedy of the commons”)
Adult workers and mother queen selected to prevent production of excess queens (“policing”)
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Comparative predictions hold
Self determination20% queen production
stingless bees
Policing of caste fate0.02% queen production
honey bees
Individual Freedom Causes a Cost to Society
But females prefer to become
queen with probability
of 56% !
Efficient Society but
No Individual Freedom
THE SAME TENSION OCCURS IN HUMAN SOCIETY !
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General conclusions Part I : empirical
– Does Wolbachia occur in ant societies? YES, IN HIGH FREQUENCY
– Alternative explanation for female biased sex-ratios in this group? PROBABLY NOT
– Other effects? INCOMPATIBILITY (SPECIATION?)
Part II : theoretical– What do animal and genomic conflicts have in
common? SAME LOGIC– Can sociobiological theory be applied to both?
YES (GENERALISED HAMILTOM’S RULE)– What do we learn from this more generally?
DEEPER INSIGHT INTO THE FUNCTIONING OF HUMAN SOCIETIES (TOC)
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The End
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AcknowledgementsProf. Dr. J. Billen Prof. Dr. R. Huybrechts Prof. Dr. J.J. Boomsma Dr. F. ItoDr. K.R. Foster Dr. F.L.W. RatnieksProf. S.A. Frank Dr. L. Sundström Dr. D.A. Grasso Drs. S. Van Borm Prof. Dr. F. Volckaert Academy of Finland, British Council,
FWO-Vlaanderen, Vlaamse Leergangen, EU Network “Social Evolution”