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Making the case for

Viral niche construction

Steven Hamblin and Mark Tanaka
University of New South Wales
... “the phenomenon of constructing, destroying, or
altering one’s environment and thus changing the
selection pressures exerted by that environment.”
Boni & Feldman, 2005
Et

Natural selection

Gene
pool

time

Genetic inheritance

t

Populations
of diverse
phenotypes

E

t+1 t+1

Natural selection

Gene
pool

Populations
of diverse
phenotypes
Et

E

t+1 t+1

Gene
pool

Populations
of diverse
phenotypes

Genetic inheritance

time

Ecological inheritance

t

Niche construction
Natural selection

Niche construction
Natural selection

Gene
pool

Populations
of diverse
phenotypes
Membracidae

Plants

Periophthalmus
Archaebacteria
Tachyoryctes
Grasshoppers
Parus
Uropeltidae Epiphytes
Characiformes Uloborus

Amphisbaenia

Prairie dogs

Typhlopidae

Finch

Tuna Algae

Hominids

Snakes
Guppies

Stickelback

Earwigs Dipterans
Parasitoid insects

Bacteria
Lycosidae

Ostariophysi

Fungi
Wasps

Chaparral

Paelobatidae

Apodidae
Jerboas Diatoms

Badgers Teiidae Gymnopphiona Lepidoptera Lanternfishes Terrapins

Carp

Ants

Scolytidae Capitonidae Vipes
Vombatidae Foxes
DermapteraRats Gerbils

Trionychidae Ant lions
Sharks
Whales Meropidae
Lovebirds

Scombridae Hamsters
Euchondrus Petrel
Corvids Beech trees Cod
Barbets Viruses

Flies

Mites

Caecilians

Monarch butterfly

Helicopsychidae

Bats

Hemisotidae

Deinopidae Apinae

Paradise fish

Beeflies

Caelifera

Hydropsychiade Lampreys

Lizards
Picidae

Skinks Sphagnum
Smelts

Reptiles
Sawflies

Hedgehogs Siluriformes
Tenebrionidae

Macrotermitinae

Bees

Rhinophrynidae Trichoptera
Aniliidae Animalia
Areneus
EnsiferanHemiptera Rabbits
Clupeiformes Eagles
Cuckoo
Ctenizidae Tuatara Apicotermes
Cicadas
Ichneumonidae Locusts
Wombats
Microhylidae Segestria
Neanderthals
Eumenidae Myrmeleontidae Ploceidae
Lutra
Scarabaeidae Pythons

Turtles

Cricitelus

Mallos Crocodiles
Mosquitoes Cassava

Fishes

Gasterosteus Weaverbirds

Flowering plants

Swifts

Alcedinidae

Momotidae

Teleutomyrmex

Bathyergidae Blattodea Hystrix
Batrachoididae Bee eaters
Motmots Tephritid flies
Rhynchocephalia Zooplankton
Opossum Snapping shrimps
Salmon Cormorants
Wren Weeverfishes
Gryllotalpidae
Leptotyphlopidae
Gophers
Moths
Mouthbrooder

Orthoptera

Lemmings

Mammals

Zebras
Crustaceans Chough

Pine trees

Alligators

Testudinata Polistinae

Squirrels

Mole rats Anomalepididae

Oviparous insects

Basidiomycetes

Eucalyptus

Pipesnakes

Birds

Beetles

Hymenoptera

Isopods

Spiders Insects
Termites

Ptilonorhynchidae

Anguillidae

Beavers

Amphipods Primates

Cyclosa

Scincidae
Galaxids

Gecko

Cyanobacteria

Osmeridae Symphyta

Vertebrates Nile monitor Passalidae

Sandpipers

Amphibians

Swallows

Ophichthidae

Homo

Arachnids

Viceroy

Cattle

Trees

Bombyliidae

Rodents
Trochilidae

Agapornis

Worms

Sea turtle
Bowerbirds

IguanasCows
Turkeys
Puffinus
Braunsapis
Cavia
Dipodidae
Bembix
Turdus
Embioptera Thrush
Woodpeckers
Eider duck
Megapodes
Coral Red admiral
Treehoppers Ariadna
Voles Tits Gastercantha Lobsters Chiromantis
Guinea pigs Vespinae
Ovenbirds Malaria
Dragonfishes
Snails Bucerotidae Chameleons
Painted lady Pisauridae
Dendrobatidae
Milkweed Magpies
Philopotamidae
House martin Acacia Cynomys Macronema Owls
Reithrodontomys Hirundines Peromyscus
Caterpillars Eels
Trachinoidei Gazelles
Theridion
Wildebeest Hirundinidae
Sardines Segestriidae Galliwasps Furnariidae Thysanoptera River otters
Liphistiidae
Limnephilidae
Storks Vanessa
Rooks Microphytes
Merganser Synbranchiformes
Coniosporium Katydids
Erinaceus Flatworms
Sarcophagids
Dolphins Chiroptera Silverfish Acrididae Catfish
Porcupines Eublepharidae Helodermatidae
Toadfish Butterflies
Stomiiformes
Tortoise Moles
Hummingbirds Snake eels
Kingfishers
Drosophila
Psocoptera
Vulpes
Braconidae Protists
Cichlids
Ergot fungus Spalax
Bark lice
Petromyzontiformes
Coleoptera
Muskrats Melinae
Atta Pleurodira
Lichens Argyroneta Honey eater Pongidae Grasses
Eubacteria Buffalo grass Tetragnatha MyctophidaeGobies Congridae Millipedes Pygopodids
Hornbills
Tapinillus Marmots
Testudinidae Mudskippers
Louse
Sauria
Mosses
Black rush Cyclorana

Dibamids

Frogs
Scombridae Hamsters
Euchondrus Petrel
Corvids Beech trees Cod
Barbets Viruses

Beeflies

Ants

Mites

Caecilians

IguanasCows

Monarch butterfly

Skinks Sphagnum
Smelts

Carp

Trionychidae Ant lions
Flies
Meropidae
Sharks
Whales
Lovebirds
Rodents
Trochilidae

Lutra

Helicopsychidae

Scolytidae Capitonidae Vipes
Vombatidae Fo
Bats
DermapteraRats Gerbils
Hemisotidae

Ploceidae

B

Paradise fish

Flowering plants

leontidae

Deinopidae Apinae

Lizards
Picidae

mbats

Hominids

Earwigs Dipterans
Parasitoid insects

Badgers Teiidae Gymnopphiona Lepidoptera Lanternfishes Terrapins

Tenebrionidae

Fishes
Neanderthals

custs

Prairie dogs

Cricitelus

Testudina

Tuatara Apicotermes

Jerboas Diatoms

Caelifera Lampreys
Hydropsychiade

Hedgehogs Siluriformes

Cuckoo

Bee

gles

Macrotermitinae

alia Areneus
era Rabbits

Lemm

Trichoptera

O

B

nobacteria

Birds

Iso

Homo

Arachnids

Amphipods Primates

hichthidae

optera

Cattle

Pipesnakes

Bowerbirds

Agapornis

Puffinus
Dipodidae
Bembix
Turdus
Embioptera Thrush
E
Megapodes
Coral Red admiral
Treehoppers Ariadna
Voles Tits Gastercantha Lobsters Chiromantis
Guinea pigs Vespinae
Snails Bucerotidae Chameleons
Painted lady Pisauridae
Dendrobatidae
Philopotamidae
Peromyscus House martin Acacia Cynomys Macronema Owls
Theridion Caterpillars Eels
Wildebeest Hirundinidae Trachinoidei Gazelles
Sardines Segestriidae Galliwasps Furnariidae Thysanoptera River otters
histiidae
Storks Vanessa
Rooks Microphytes
Merganser Synbranchiforme
Coniosporium Katydids
Sarcophagids Erinaceus Flatworms
era Silverfish Acrididae Catfish
Porcupines Eublepharidae He
oadfish Butterflies
Tortoise Moles
Hummingbirds Snake eels
shers
Psocoptera
Vulpes Drosophila
Braconidae Pro
Cichlids
ungus Spalax
Petromyzontiformes
Melinae Bark lice
Coleoptera
Muskrats
Atta Pleurodira
Lichens Argyroneta Honey eater Pongidae Grasses
Eubacteria Buffalo grass Tetragnatha MyctophidaeGobies Congridae Millipedes Pygopodids
Hornbills
us Marmots
Testudinidae Mudskip
Louse
a
Mosses
Black rush Cyclorana
Cavia

Dibamids
2 Gt/y

5 Gt/y

Atmosphere
805 Gt
+4 Gt/y
Biological and chemical
processes

Fossil fuel use

Deforestation

CO2

UV
CO2 90 Gt/y

93 Gt/y

DOC
700 Gt
Plankton
5 Gt

Fossil fuels
7, 000 Gt
Thermocline

Ocean
38,500 Gt

POC
50 Gt

Plankton infected
by virus
Lysis of plankton
by viruses
150 Gt/y
Uninfected cells and
detritus sink
3 Gt/y

Figure 4 | Viruses can affect the efficiency of the biological pump. Viruses cause the lysis of cells, converting them into particulate organic carbon (POC) and
dissolved organic carbon (DOC). This reduces the rate atwhich C sinks from the surface layer into the deep ocean where the carbon is trapped for millennia
(biological pump). Instead the carbon is retained in the surface waters where it is photo-oxidized and respired, in chemical equilibrium with the atmosphere.
The net effect is a faster rate of CO2 build-up in the atmosphere than would occur if the POC were ‘exported’ to the deep ocean.

oligotrophic ocean, where slow rate processes make it difficult to
obtain reliable data except from visibly infected cells or viral decay
rates. Moreover, the assumptions for calculating mortality rates are
particularly poorly grounded for open-ocean species. Even within relatively productive environments, estimates of the contribution of

ton, where virus-infected cells sink rapidly78, potentially increasing the
transport of cells to deeper waters27. Nutrients other than carbon are
also released by viral lysis79,80. As these nutrients are largely organically
bound, this can affect their availability and pathways of cycling. In
some cases, released nutrients such as iron can fill a major portion of

Suttle, 2005
Viral niche construction!
Case study:

Baculoviridae

http://www.mardre.com/homepage/mic/tem/samples/bio/virus/bac2.htm
Nucleopolyhedrovirus
(NPV)
MNPV

Virions

Occlusion
body
Granulovirus
(GV)

SNPV

Figure 1 Occluded virions. Baculovirus genera are based on the structures
of occluded virus, polyhedral shaped for nucleopolyhedrovirus (NPV) with
many enveloped nucleocapsids or smaller ovoid occlusions with a single
nucleocapsid for granulovirus (GV). NPVs may contain multiple
nucleocapsids (MNPV) or single nucleocapsids (SNPV).

genera should be revised. The lepidopteran NPVs are more

the point where they a
have lost the genes enti
that the hymenopteran
midgut, in which case a
necessary (Duffy et al
Gammabaculoviridae is
drovirus (NeleNPV).
Only one member of
sequenced to date. Cule
(CuniNPV) infects mos
produces both BV (con
matrix protein shows no
or granulin and may r
proteins (Perera et al.,
Most baculoviruses
are named after the i
exception to this rule is
32 different lepidoptera

Structure

Guarino, 2011
Lymantria dispar
Zombie / Gooey
Zombie / Gooey
egt

chit
cath

Hawtin et al, 1997
Hoover et al, 2011
Zombie / Gooey

How do you acquire both features?
Zombie but
not gooey?
Non-zombie, but gooey?
µ
Exposed

Susceptible

⌫

Carrier

Liquefied

⌧
L+

0

C

✓

Reservoir

✏
R0 =

⇣

0

+ +✓

⌘⇣

⌫
⌫+µ

⌘

+

⇣

+⌧

⌘⇣

⌫
⌫+µ

⌘⇣

+ +✓

⌘
R0 =

⇣

0

+ +✓

⌘⇣

⌫
⌫+µ

⌘

+

⇣

+⌧

⌘⇣

⌫
⌫+µ

⌘⇣

+ +✓

µ
Exposed

Susceptible

⌫

Carrier

Liquefied

⌧
L+

0

C

✓

Reservoir

✏

⌘
R0 =

⇣

0

+ +✓

⌘⇣

⌫
⌫+µ

⌘

+

⇣

+⌧

⌘⇣

⌫
⌫+µ

⌘⇣

+ +✓

µ
Exposed

Susceptible

⌫

Carrier

Liquefied

⌧
L+

0

C

✓

Reservoir

✏

⌘
R0 =

⇣

0

+ +✓

⌘⇣

⌫
⌫+µ

⌘

+

⇣

+⌧

⌘⇣

⌫
⌫+µ

⌘⇣

+ +✓

µ
Exposed

Susceptible

⌫

Carrier

Liquefied

⌧
L+

0

C

✓

Reservoir

✏

⌘
R0 =

⇣

0

+ +✓

⌘⇣

⌫
⌫+µ

⌘

+

⇣

+⌧

⌘⇣

⌫
⌫+µ

⌘⇣

+ +✓

µ
Exposed

Susceptible

⌫

Carrier

Liquefied

⌧
L+

0

C

✓

Reservoir

✏

⌘
R0 =

⇣

0

+ +✓

⌘⇣

⌫
⌫+µ

⌘

+

⇣

+⌧

⌘⇣

⌫
⌫+µ

⌘⇣

+ +✓

µ
Exposed

Susceptible

⌫

Carrier

Liquefied

⌧
L+

0

C

✓

Reservoir

✏

⌘
R0 =

⇣

0

+ +✓

⌘⇣

⌫
⌫+µ

⌘

+

⇣

+⌧

⌘⇣

⌫
⌫+µ

⌘⇣

+ +✓

µ
Exposed

Susceptible

⌫

Carrier

Liquefied

⌧
L+

0

C

✓

Reservoir

✏

⌘
Genotypes
⌧
ZG
Zg
zG
zg

(1 + )

⌧ (1

(1

⌧
⌧ (1 + )
⌧

)

)
0
0

✓
✓L
✓L
✓H
✓H

Zombie / Gooey

Genotypes and their e↵ect on the model para

z

g
, ⌧, 0, ✓H

(1

G
), ⌧ (1 + ), , ✓H
So...

niche construction?
1.0

% of population

Susceptible

0.8

Exposed
Cadaver

0.6

Liquefied
Reservoir

0.4

0.2

0.0

20

40

60

80

Time (days)

100

120

140

t
25

20

15

Genotype

R0

zg
zG
Zg
ZG

10

5

0
0.0

0.2

0.4

γ

0.6

0.8
25

20

15

Genotype

R0

zg
zG
Zg
ZG

10

5

0
0.0

0.2

0.4

γ

0.6

0.8
25

Fitness sequence:

20

zg < Zg < zG < ZG
Genotype
zg
zG
Zg
ZG

R0

=

15

10

5

0
0.0

0.2

0.4

γ

0.6

0.8
0.5

0.4

Sequence

n

θ

(reservoir contributio
by carriers)

No NC
Zg,zG,zg
zG,Zg,zg
Zg,zg,zG
zG,zg,Zg
zg,Zg,zG
zg,zG,Zg

0.3

0.2

0.1

0.5

1.0

1.5

2.0

β (transmission from liquefied hosts)
egt
chit
cath
null

Culex nigripalpus NPV
Neodiprion abietis NPV
Neodiprion lecontii NPV
Neodiprion sertifer NPV
Pieris rapae GV
Choristoneura occidentalis GV
Clostera anachoreta GV
Cryptophlebia leucotreta GV
Cydia pomonella GV
Adoxophyes orana GV
Phthorimaea operculella GV
Agrotis segetum GV
Plutella xylostella GV
Spodoptera litura GV
Pseudaletia unipuncta GV
Helicoverpa armigera GV
Xestia c-nigrum GV
Leucania separata NPV
Spodoptera litura NPV
Choristoneura fumiferana MNPV
Orgyia pseudotsugata MNPV
Hyphantria cunea NPV
Antheraea pernyi NPV
Epiphyas postvittana NPV
Anticarsia gemmatalis NPV
Choristoneura fumiferana DEF MNPV
Maruca vitrata MNPV
Bombyx mandarina NPV
Bombyx mori NPV
Plutella xylostella multiple NPV
Rachiplusia ou MNPV
Autographa californica NPV
Lymantria xylina MNPV
Lymantria dispar MNPV
Helicoverpa armigera NPV NNg1
Helicoverpa zea SNPV
Helicoverpa armigera NPV
Helicoverpa armigera NPV G4
Adoxophyes orana NPV
Adoxophyes honmai NPV
Clanis bilineata NPV virus
Ecotropis obliqua NPV
Euproctis pseudoconspersa NPV
Orgyia leucostigma NPV
Trichoplusia ni SNPV
Chrysodeixis chalcites NPV
Helicoverpa armigera multiple NPV
Mamestra configurata NPV-B
Mamestra configurata NPV-A
Agrotis ipsilon multiple NPV
Agrotis segetum NPV
Spodoptera frugiperda MNPV
Spodoptera litura NPV II
Spodoptera exigua MNPV
Granulovirus Morphs

Type I

•Infect only midgut and fat bodies.
•Kill slowly, little or no liquefaction.

Type II

•Infect multiple tissues.
•Kill more quickly, liquefy the host.
Pieris rapae GV

Zombie/Gooey

egt

Choristoneura occidentalis GV

chit
cath

Clostera anachoreta GV

Cryptophlebia leucotreta GV

Cydia pomonella GV

Adoxophyes orana GV

Phthorimaea operculella GV

Agrotis segetum GV

Plutella xylostella GV

Spodoptera litura GV

egt
chit
cath
null

Type I
Type II

Pseudaletia unipuncta GV

Helicoverpa armigera GV

Xestia c-nigrum GV
So...

niche construction!
Viruses are driving behaviour
... for their own benefit.
Sociality?
Biopesticides
for fun and profit!
Thanks!
Tanaka Lab
Photos
Kelli Hoover used by permission
,

ratsinis (Flickr) used under a CC license

GollyGForce (Flickr) used under a CC license

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ISBE 2012

  • 1. Making the case for Viral niche construction Steven Hamblin and Mark Tanaka University of New South Wales
  • 2. ... “the phenomenon of constructing, destroying, or altering one’s environment and thus changing the selection pressures exerted by that environment.” Boni & Feldman, 2005
  • 3. Et Natural selection Gene pool time Genetic inheritance t Populations of diverse phenotypes E t+1 t+1 Natural selection Gene pool Populations of diverse phenotypes
  • 4. Et E t+1 t+1 Gene pool Populations of diverse phenotypes Genetic inheritance time Ecological inheritance t Niche construction Natural selection Niche construction Natural selection Gene pool Populations of diverse phenotypes
  • 5. Membracidae Plants Periophthalmus Archaebacteria Tachyoryctes Grasshoppers Parus Uropeltidae Epiphytes Characiformes Uloborus Amphisbaenia Prairie dogs Typhlopidae Finch Tuna Algae Hominids Snakes Guppies Stickelback Earwigs Dipterans Parasitoid insects Bacteria Lycosidae Ostariophysi Fungi Wasps Chaparral Paelobatidae Apodidae Jerboas Diatoms Badgers Teiidae Gymnopphiona Lepidoptera Lanternfishes Terrapins Carp Ants Scolytidae Capitonidae Vipes Vombatidae Foxes DermapteraRats Gerbils Trionychidae Ant lions Sharks Whales Meropidae Lovebirds Scombridae Hamsters Euchondrus Petrel Corvids Beech trees Cod Barbets Viruses Flies Mites Caecilians Monarch butterfly Helicopsychidae Bats Hemisotidae Deinopidae Apinae Paradise fish Beeflies Caelifera Hydropsychiade Lampreys Lizards Picidae Skinks Sphagnum Smelts Reptiles Sawflies Hedgehogs Siluriformes Tenebrionidae Macrotermitinae Bees Rhinophrynidae Trichoptera Aniliidae Animalia Areneus EnsiferanHemiptera Rabbits Clupeiformes Eagles Cuckoo Ctenizidae Tuatara Apicotermes Cicadas Ichneumonidae Locusts Wombats Microhylidae Segestria Neanderthals Eumenidae Myrmeleontidae Ploceidae Lutra Scarabaeidae Pythons Turtles Cricitelus Mallos Crocodiles Mosquitoes Cassava Fishes Gasterosteus Weaverbirds Flowering plants Swifts Alcedinidae Momotidae Teleutomyrmex Bathyergidae Blattodea Hystrix Batrachoididae Bee eaters Motmots Tephritid flies Rhynchocephalia Zooplankton Opossum Snapping shrimps Salmon Cormorants Wren Weeverfishes Gryllotalpidae Leptotyphlopidae Gophers Moths Mouthbrooder Orthoptera Lemmings Mammals Zebras Crustaceans Chough Pine trees Alligators Testudinata Polistinae Squirrels Mole rats Anomalepididae Oviparous insects Basidiomycetes Eucalyptus Pipesnakes Birds Beetles Hymenoptera Isopods Spiders Insects Termites Ptilonorhynchidae Anguillidae Beavers Amphipods Primates Cyclosa Scincidae Galaxids Gecko Cyanobacteria Osmeridae Symphyta Vertebrates Nile monitor Passalidae Sandpipers Amphibians Swallows Ophichthidae Homo Arachnids Viceroy Cattle Trees Bombyliidae Rodents Trochilidae Agapornis Worms Sea turtle Bowerbirds IguanasCows Turkeys Puffinus Braunsapis Cavia Dipodidae Bembix Turdus Embioptera Thrush Woodpeckers Eider duck Megapodes Coral Red admiral Treehoppers Ariadna Voles Tits Gastercantha Lobsters Chiromantis Guinea pigs Vespinae Ovenbirds Malaria Dragonfishes Snails Bucerotidae Chameleons Painted lady Pisauridae Dendrobatidae Milkweed Magpies Philopotamidae House martin Acacia Cynomys Macronema Owls Reithrodontomys Hirundines Peromyscus Caterpillars Eels Trachinoidei Gazelles Theridion Wildebeest Hirundinidae Sardines Segestriidae Galliwasps Furnariidae Thysanoptera River otters Liphistiidae Limnephilidae Storks Vanessa Rooks Microphytes Merganser Synbranchiformes Coniosporium Katydids Erinaceus Flatworms Sarcophagids Dolphins Chiroptera Silverfish Acrididae Catfish Porcupines Eublepharidae Helodermatidae Toadfish Butterflies Stomiiformes Tortoise Moles Hummingbirds Snake eels Kingfishers Drosophila Psocoptera Vulpes Braconidae Protists Cichlids Ergot fungus Spalax Bark lice Petromyzontiformes Coleoptera Muskrats Melinae Atta Pleurodira Lichens Argyroneta Honey eater Pongidae Grasses Eubacteria Buffalo grass Tetragnatha MyctophidaeGobies Congridae Millipedes Pygopodids Hornbills Tapinillus Marmots Testudinidae Mudskippers Louse Sauria Mosses Black rush Cyclorana Dibamids Frogs
  • 6. Scombridae Hamsters Euchondrus Petrel Corvids Beech trees Cod Barbets Viruses Beeflies Ants Mites Caecilians IguanasCows Monarch butterfly Skinks Sphagnum Smelts Carp Trionychidae Ant lions Flies Meropidae Sharks Whales Lovebirds Rodents Trochilidae Lutra Helicopsychidae Scolytidae Capitonidae Vipes Vombatidae Fo Bats DermapteraRats Gerbils Hemisotidae Ploceidae B Paradise fish Flowering plants leontidae Deinopidae Apinae Lizards Picidae mbats Hominids Earwigs Dipterans Parasitoid insects Badgers Teiidae Gymnopphiona Lepidoptera Lanternfishes Terrapins Tenebrionidae Fishes Neanderthals custs Prairie dogs Cricitelus Testudina Tuatara Apicotermes Jerboas Diatoms Caelifera Lampreys Hydropsychiade Hedgehogs Siluriformes Cuckoo Bee gles Macrotermitinae alia Areneus era Rabbits Lemm Trichoptera O B nobacteria Birds Iso Homo Arachnids Amphipods Primates hichthidae optera Cattle Pipesnakes Bowerbirds Agapornis Puffinus Dipodidae Bembix Turdus Embioptera Thrush E Megapodes Coral Red admiral Treehoppers Ariadna Voles Tits Gastercantha Lobsters Chiromantis Guinea pigs Vespinae Snails Bucerotidae Chameleons Painted lady Pisauridae Dendrobatidae Philopotamidae Peromyscus House martin Acacia Cynomys Macronema Owls Theridion Caterpillars Eels Wildebeest Hirundinidae Trachinoidei Gazelles Sardines Segestriidae Galliwasps Furnariidae Thysanoptera River otters histiidae Storks Vanessa Rooks Microphytes Merganser Synbranchiforme Coniosporium Katydids Sarcophagids Erinaceus Flatworms era Silverfish Acrididae Catfish Porcupines Eublepharidae He oadfish Butterflies Tortoise Moles Hummingbirds Snake eels shers Psocoptera Vulpes Drosophila Braconidae Pro Cichlids ungus Spalax Petromyzontiformes Melinae Bark lice Coleoptera Muskrats Atta Pleurodira Lichens Argyroneta Honey eater Pongidae Grasses Eubacteria Buffalo grass Tetragnatha MyctophidaeGobies Congridae Millipedes Pygopodids Hornbills us Marmots Testudinidae Mudskip Louse a Mosses Black rush Cyclorana Cavia Dibamids
  • 7.
  • 8.
  • 9. 2 Gt/y 5 Gt/y Atmosphere 805 Gt +4 Gt/y Biological and chemical processes Fossil fuel use Deforestation CO2 UV CO2 90 Gt/y 93 Gt/y DOC 700 Gt Plankton 5 Gt Fossil fuels 7, 000 Gt Thermocline Ocean 38,500 Gt POC 50 Gt Plankton infected by virus Lysis of plankton by viruses 150 Gt/y Uninfected cells and detritus sink 3 Gt/y Figure 4 | Viruses can affect the efficiency of the biological pump. Viruses cause the lysis of cells, converting them into particulate organic carbon (POC) and dissolved organic carbon (DOC). This reduces the rate atwhich C sinks from the surface layer into the deep ocean where the carbon is trapped for millennia (biological pump). Instead the carbon is retained in the surface waters where it is photo-oxidized and respired, in chemical equilibrium with the atmosphere. The net effect is a faster rate of CO2 build-up in the atmosphere than would occur if the POC were ‘exported’ to the deep ocean. oligotrophic ocean, where slow rate processes make it difficult to obtain reliable data except from visibly infected cells or viral decay rates. Moreover, the assumptions for calculating mortality rates are particularly poorly grounded for open-ocean species. Even within relatively productive environments, estimates of the contribution of ton, where virus-infected cells sink rapidly78, potentially increasing the transport of cells to deeper waters27. Nutrients other than carbon are also released by viral lysis79,80. As these nutrients are largely organically bound, this can affect their availability and pathways of cycling. In some cases, released nutrients such as iron can fill a major portion of Suttle, 2005
  • 12. Nucleopolyhedrovirus (NPV) MNPV Virions Occlusion body Granulovirus (GV) SNPV Figure 1 Occluded virions. Baculovirus genera are based on the structures of occluded virus, polyhedral shaped for nucleopolyhedrovirus (NPV) with many enveloped nucleocapsids or smaller ovoid occlusions with a single nucleocapsid for granulovirus (GV). NPVs may contain multiple nucleocapsids (MNPV) or single nucleocapsids (SNPV). genera should be revised. The lepidopteran NPVs are more the point where they a have lost the genes enti that the hymenopteran midgut, in which case a necessary (Duffy et al Gammabaculoviridae is drovirus (NeleNPV). Only one member of sequenced to date. Cule (CuniNPV) infects mos produces both BV (con matrix protein shows no or granulin and may r proteins (Perera et al., Most baculoviruses are named after the i exception to this rule is 32 different lepidoptera Structure Guarino, 2011
  • 15. Zombie / Gooey egt chit cath Hawtin et al, 1997 Hoover et al, 2011
  • 16. Zombie / Gooey How do you acquire both features?
  • 21. R0 = ⇣ 0 + +✓ ⌘⇣ ⌫ ⌫+µ ⌘ + ⇣ +⌧ ⌘⇣ ⌫ ⌫+µ ⌘⇣ + +✓ µ Exposed Susceptible ⌫ Carrier Liquefied ⌧ L+ 0 C ✓ Reservoir ✏ ⌘
  • 22. R0 = ⇣ 0 + +✓ ⌘⇣ ⌫ ⌫+µ ⌘ + ⇣ +⌧ ⌘⇣ ⌫ ⌫+µ ⌘⇣ + +✓ µ Exposed Susceptible ⌫ Carrier Liquefied ⌧ L+ 0 C ✓ Reservoir ✏ ⌘
  • 23. R0 = ⇣ 0 + +✓ ⌘⇣ ⌫ ⌫+µ ⌘ + ⇣ +⌧ ⌘⇣ ⌫ ⌫+µ ⌘⇣ + +✓ µ Exposed Susceptible ⌫ Carrier Liquefied ⌧ L+ 0 C ✓ Reservoir ✏ ⌘
  • 24. R0 = ⇣ 0 + +✓ ⌘⇣ ⌫ ⌫+µ ⌘ + ⇣ +⌧ ⌘⇣ ⌫ ⌫+µ ⌘⇣ + +✓ µ Exposed Susceptible ⌫ Carrier Liquefied ⌧ L+ 0 C ✓ Reservoir ✏ ⌘
  • 25. R0 = ⇣ 0 + +✓ ⌘⇣ ⌫ ⌫+µ ⌘ + ⇣ +⌧ ⌘⇣ ⌫ ⌫+µ ⌘⇣ + +✓ µ Exposed Susceptible ⌫ Carrier Liquefied ⌧ L+ 0 C ✓ Reservoir ✏ ⌘
  • 26. R0 = ⇣ 0 + +✓ ⌘⇣ ⌫ ⌫+µ ⌘ + ⇣ +⌧ ⌘⇣ ⌫ ⌫+µ ⌘⇣ + +✓ µ Exposed Susceptible ⌫ Carrier Liquefied ⌧ L+ 0 C ✓ Reservoir ✏ ⌘
  • 27. Genotypes ⌧ ZG Zg zG zg (1 + ) ⌧ (1 (1 ⌧ ⌧ (1 + ) ⌧ ) ) 0 0 ✓ ✓L ✓L ✓H ✓H Zombie / Gooey Genotypes and their e↵ect on the model para z g , ⌧, 0, ✓H (1 G ), ⌧ (1 + ), , ✓H
  • 32. 25 Fitness sequence: 20 zg < Zg < zG < ZG Genotype zg zG Zg ZG R0 = 15 10 5 0 0.0 0.2 0.4 γ 0.6 0.8
  • 33. 0.5 0.4 Sequence n θ (reservoir contributio by carriers) No NC Zg,zG,zg zG,Zg,zg Zg,zg,zG zG,zg,Zg zg,Zg,zG zg,zG,Zg 0.3 0.2 0.1 0.5 1.0 1.5 2.0 β (transmission from liquefied hosts)
  • 34. egt chit cath null Culex nigripalpus NPV Neodiprion abietis NPV Neodiprion lecontii NPV Neodiprion sertifer NPV Pieris rapae GV Choristoneura occidentalis GV Clostera anachoreta GV Cryptophlebia leucotreta GV Cydia pomonella GV Adoxophyes orana GV Phthorimaea operculella GV Agrotis segetum GV Plutella xylostella GV Spodoptera litura GV Pseudaletia unipuncta GV Helicoverpa armigera GV Xestia c-nigrum GV Leucania separata NPV Spodoptera litura NPV Choristoneura fumiferana MNPV Orgyia pseudotsugata MNPV Hyphantria cunea NPV Antheraea pernyi NPV Epiphyas postvittana NPV Anticarsia gemmatalis NPV Choristoneura fumiferana DEF MNPV Maruca vitrata MNPV Bombyx mandarina NPV Bombyx mori NPV Plutella xylostella multiple NPV Rachiplusia ou MNPV Autographa californica NPV Lymantria xylina MNPV Lymantria dispar MNPV Helicoverpa armigera NPV NNg1 Helicoverpa zea SNPV Helicoverpa armigera NPV Helicoverpa armigera NPV G4 Adoxophyes orana NPV Adoxophyes honmai NPV Clanis bilineata NPV virus Ecotropis obliqua NPV Euproctis pseudoconspersa NPV Orgyia leucostigma NPV Trichoplusia ni SNPV Chrysodeixis chalcites NPV Helicoverpa armigera multiple NPV Mamestra configurata NPV-B Mamestra configurata NPV-A Agrotis ipsilon multiple NPV Agrotis segetum NPV Spodoptera frugiperda MNPV Spodoptera litura NPV II Spodoptera exigua MNPV
  • 35. Granulovirus Morphs Type I •Infect only midgut and fat bodies. •Kill slowly, little or no liquefaction. Type II •Infect multiple tissues. •Kill more quickly, liquefy the host.
  • 36. Pieris rapae GV Zombie/Gooey egt Choristoneura occidentalis GV chit cath Clostera anachoreta GV Cryptophlebia leucotreta GV Cydia pomonella GV Adoxophyes orana GV Phthorimaea operculella GV Agrotis segetum GV Plutella xylostella GV Spodoptera litura GV egt chit cath null Type I Type II Pseudaletia unipuncta GV Helicoverpa armigera GV Xestia c-nigrum GV
  • 38. Viruses are driving behaviour ... for their own benefit.
  • 42. Photos Kelli Hoover used by permission , ratsinis (Flickr) used under a CC license GollyGForce (Flickr) used under a CC license