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EVOLUTION OF
RESOURCE HARVESTING
ORGANS
WITH RESILIENCE OF INSECTS
AND ROLE OF PLANTS
FOR SUSTENANCE OF INSECT
DIVERSITY AND MODES OF
INSECT PLANT INTERACTION
Introduction
v Theinsecta are by far the most species-richtaxon.
v Several hypotheses have been propounded to
explain the evolution of the striking diversity of
the Insecta and in particular of herbivorous
insects.
vAnew insect taxon could become established only
if another was excluded by competition and
becameextinct.
Evolutionary diversity of insect ears
•Insects are among the oldest land animals, and exist for
more than 400 millionyears.
•Among insects, tympanal earsevolved at least
18 times, resulting in adiversity of auditory systems.
•Insects use their ears in different behavioural contexts,
mainly intraspecific communication for mate attraction,
predator avoidance, and parasitic host localisation.
Evolutionary origin of auditory behaviour
v It is driven by natural and sexual selection, insect ears
evolved mainly for intraspecific communication and
predator detection.
v The most prominent example is between bats and
nocturnal flying insects.
v Ears of bats should be highly sensitive to ultrasound.
Hemiptera of small body size use ancestrally tymbal
mechanisms to generate vibration signals and Cicadidae
produce auditory sounds by identical means of tymbal
mechanisms.
Ten regions where tympanal
organs have evolved within
insect groups
1. Lepidoptera; Sphingidae
2. Orthoptera; Ensifera
3. Diptera; Tachinidae
4. Mantodea; Mantidae
5. Lepidoptera;
Geometroidea and
Pyraloidea
6. Orthoptera; Acrididae
7. Hemiptera; Cicadidae
8. Lepidoptera; Noctuoidea
9. Hemiptera; Corixidae
10. Neuroptera; Chrysopidae
Evolutionary origin of insect wings
Theories of wing origin:
1.Paranotal theory
2.Epicoxal theory
3.Endite- Exite theory
Epicoxal Hypothesis-
Wings from abdominal gills
But lacking the tarsus, where the wing’s coastal
surface normally would be.
Role of Plants for sustenance of insect diversity
Various factors have been proposed as agents of
selection in the evolution of the specialistic feeding
habit of herbivorous insects.
Themain factors are:
(1) coping with plant secondarymetabolites
(2) avoiding competition
(3) reducing mortality fromnatural enemies.
Coping with plant secondary metabolites
• Insects that are capable of detoxifying one class of plant
compound usually cannot detoxify a very different class of
secondary metabolites.
• Specialist insects using the same plant taxon have evolved
different detoxification or excretion mechanisms to avoid the
impact of the samesecondary plantmetabolites.
• These are steroidal compounds responsible for moulting, growth
and maturation of insects.
• It is produced by prothoracicgland.
• An exciting investigation has shown that a single plant gene can
determine whether the plant will be included in the diet of insect
herbivores.
Herbivory
v Herbivory (phytophagy): Leafchewing , sapsacking,
seedpredation, gall inducing, leafmining
v Insect-plant mutualism –pollination and plantinsect
food for defence relationshipsHerbivory
v Chewersmost diverse of theleaf chewing insects are
the Coleoptera and Lepidoptera.
v Insects eat leaves,roots, shoots, stems, andflowers or
fruits
v Chewing insects possessmandibulate mouthparts
v Mandibles serve to cut and grindfood
v Mandibles are highly sclerotized to reducewear
v High silica content and cellulose canact asresistance to
herbivory
Mining and boring
• Insects live in between 2 epidermal layers of aleaf.
Damageappears astunnels , blotches or blisters.
• Independently evolved in 4 orders : Diptera,Lepidoptera,
Coleoptera, and Hymenoptera
• Different speciesmay excavate different layers of leaf
parenchyma or reside in particularleaf
• Fruit boring
• Stem boring
• Wood borers
• Stalk boring
• Plant boring
Mining and boring
Sap sucking
• Drains plants resources by tapping into xylem and
phloem
• Hemipterans exemplify this strategy through
haustellate mouth parts and Serves to pierce
tissues and suckliquid food.
• Labium modified into asheath enclosingstylet
maxillae
• Stylets pierce cuticle and canchangeorientation
• Foodchannels empties into cibarialcavity
Sap sucking
Chewers
• No relative sizerestrictions
• Heavymechanical damage
• Facedwith indigestible compounds andtoxins
Suckers
• Restricted to arelatively smallsize
• Avoid mechanical damage (but still damaging)
• Avoid indigestible compounds and mosttoxins
• Xylem less suitable
Gall makers
• Gallsconsists of pathologically developed cells, tissues
or organs of plants that have arisen by hypertrophy
and/or hyperplasia asa result of stimulation from
foreign organisms.
• Orders that makesgalls;Hemiptera Diptera
Hymenoptera
Pollination
• A pollinator
moves pollen
is an animal that
from the male anther of
aflower to the female stigma ofaflower.
• This helps to bring about fertilization of the
ovules in the flower by the male gametes
from the pollengrains.
• Apollinator is different from apollenizer.
• A plant that is a source of pollen for the
pollination process.
Types of pollinators
Bees
• Themost recognized pollinators are the
various speciesof bees, which areplainly
adapted to pollination.
• Beestypically are fuzzy andcarry
an electrostatic charge.
• Both features help pollen grains adhere to
their bodies, but theyalso have specialized
pollen-carrying structures
Honey bees
• Honey bees travel from flower to flower, collecting
nectar (later converted tohoney), and pollen grains.
• The bee collects the pollen by rubbing against the
anthers.
• Nectar provides the energy for bee nutrition; pollen
provides the protein.
• Good pollination management seeks to have bees in
a "building" state during the bloom period of the
crop, thus requiring them to
gather pollen, and making them
more efficientpollinators.
Other insects
• Many insects other than beesaccomplish pollination by
visiting flowers for nectar or pollen, or commonly both.
• Many do so adventitiously, but the most important
pollinators are specialists for at least parts of their
lifecycles for at least certainfunctions.
• Many bee flies, and someTabanidaeand Nemestrinidae
are particularly adapted to pollinating fynbos and
Karoo plants with narrow, deep corolla tubes, such as
Lapeirousia species.
Predation
• Predation is a biological interaction where a predator feeds on its
prey.
• An insect predator is large in size, active in habits and have
structural adoptions for catching the prey with well
development senceorgans and capacity for swiftmovements.
• Theimportant entomophagous predators are:
Ø Dragonflies(Odonata): Thenaiads feed on aquatic insects and the
adult on insects like mosquitoes, flies and moths while on the
wings.
Ø Praying mantids(Dictyoptera): feed upon flies, grasshopper and
caterpillers.
Ø Lady bird beetle(Coccinellidae): feed on soft body insects.
Ø Tiger beetle( cicindellidae): Cicindella spp. feed upon avariety of
insects.
Ø Ground beetle(Carabidae): feed upon Opisina arenosella.
Predation
Parasitism
• Parasitism is anon-mutual relationship between specieswhere
one species, the parasite, benefits at the expense of the other, the
host.
• Types:Insect parasitism is of fourkinds:
a. Simple parasitism: Thisterm is applied when there is asingle attack of
the parasitoid on the host.irrespective of the number of the eggs laid.
b. Super parasitism: Parasitization of ahost by more than oneparasitic
individual usually of one kind -used especially of parasitic insects.
c. Multiple parasitism: Acondition in which parasites of differentspecies
parasitize asingle host, in contrast to superparasitism or
hyperparasitism.
d. Hyperparasitism: It meansattack of aparasitiod on an insect which.is
already aparasite asin the caseof bethylid and braconid parasites of
Opisina arenosella.
Mode of Insect Plant
Interaction
Overview
• Brief introduction: herbivory & arms race
• Arms / Weapons:
– Diversity of plant defenses
– Diversity of insect offence / counterdefense
• Signaling interactions:
– Signals that plants received from herbivores
– Signals that insects perceived from plants
Herbivory: host range
Oliogophagy
(<3 plant families )
Polyphagy
Generalist
Monophagy
Specialist
Herbivory: an arms race
Insects attack
consume plants
Plants defend
against being eaten
warfare
The arms race: coevolution
Successful
consumer
New defenses
Successful
defender
New offenses
stepwise reciprocal changes
The arms race: a typical example
Insect resistant to toxin 3
Insect resistant to toxin2
Insect resistant to toxin1
Insect eats plant
Plants make toxin 1
Plants make toxin 2
Plants make toxin 3
Plants make toxin 4
Arms: diversity of plant defenses
• Avoidance---escape from herbivores, no actual feeding.
• Resistance---reduce fitness of insects after contact
• Tolerance---to stand and take it simply by outgrowing
the damage---compensatory growth
Plant defenses: avoidance
• Escape in time
• Escape in space
• Chemical
escape
(repellent
production,
no attractant)
• Morphological
escape
• Heliconius
butterflies
avoid laying
eggs on plants
already
occupied by
eggs
• Plants
Passiflora
create fake
yellow
eggs
Plant defenses: Resistance
• Morphological resistance
– hairs, spines, hook, sticky glands,
immobilizing insects or puncturing their body
wall
• Chemical resistance
– Various toxic compounds
• Mechanical resistance
– squirt-gun
Mechanical resistance: Bursera’s
squirt-gun defense
Becerra, J. X., 2003, PNAS 100 (22), 12804-12807
Morphological resistance : thorns
• The pronounced
thorns endowed upon
the trunk of this tree
even can fend off
vertebrate herbivores,
such as sloths and
monkeys.
Chemical resistance: diversity
& classification
• Chronic (quantitative) defense: Large/complex
molecules reduce digestibility / nutrition
(tannins, lignins, cellulose, silica, etc).
• Acute (qualitative) defense: Small / simple
molecules target specific insect system.
1.Toxic amino acids
2.Toxic proteins
3. Proteinase inhibitors (=PIs)
4. Allelochemicals (=secondary comounds)
cyanide, glycosides, alkaloids, terpenoids, saponins,
flavenoids, furanocoumarins, indoles, phytoecdysteroids
Chemical resistance:
indirect defense
Plants
Herbivores
Parasitoids & Predators
Mutualism
Arms: Insect’s offense/counterdefenses
• Behavioral avoidance: selectively choose
hostplants. feeding/oviposition choice
• Modify its own behavior, or biochemistry
to overcome plant defense
• Behavioral counterdefense (or offense)
• Biochemical counterdefense
• Actively manipulate hostplants’s nutrition
& defense
Insect’s counterdefenses:
Behavioral offense
• Vein cutting
• trenching
• Leaf rolling
• Mining
• Gardening
• Gregarious feeding
Behavioral offense: vein cutting
Specialized Blepharida’s
counterdefense
vein cutting
Becerra, J. X., 2003, PNAS 100 (22), 12804-12807
Bursera’s defense
squirt-gun
Behavioral offense: trenching
avoid intoxication by
trenching the laticifers
upstream of their intended
feeding site (Wittstock &
Gershenzon, 2002)
Larvae of Erinnyis alope
starting to feed after servering
(trenching) a Carica papaya
Behavioral offense: leaf-rolling
Berenbaum & Zangerl: webworm-parsnip chemical co-evolution
Behavioral offense: gardening
l Leaf cutter ants
(Azteca) use leaf
material and flower
to culture a fungal
garden (facilitate
food storage and
toxin degradation),
which is then used
for food.
Behavioral offense :gregarious feeding
(group counterdefense)
l Sunn Pest: gregarious feeding on wheat
l increases host plant susceptibility
l Pine beetle: calling for help if needed
Insect’s counterdefenses:
Biochemical counterdefense
• Rapid excretion
• Sequestration of toxins
• Detoxification of toxins: cytochrome P450
monooxygenase (P450), esterase,
glutathione-S-transferase (GST), etc.
• Target site insensitivity
Biochemical counterdefense:
rapid excretion
Experiment:
Nicotine dose in artificial diet –> 93% excreted within 2 hours!
Experiment:
Nicotine injected into hemolymph Manduca sexta
Biochemical counterdefense:
sequestration
• Unpalatable insects sequestrate allelochemicals for their
own defense
• Many also develop warning color or/and gregarious feeding
Biochemical counterdefense:
detoxification
700
600
500
400
300
200
100
0
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Vmax/Km
Cotton bollworm P450 CYP6B8 can
metabolize both allelochemicals and
insecticides
Li, X., Berenbaum M. R, & Schuler, M.A., 2004, PNAS
Biochemical counterdefense:
target site insensitivity
Na+,K+-ATPase is sensitive to ouabain (a cardiac
glycoside). In Monarch butterflies a single-point mutation
resulted in insensitive ATPase.
Other example: induction of proteases in herbivore gut,
which are insensitive of proteinase inhibitors
Insect’s counterdefenses:
manipulate hostplants
• Manipulate plants’ growth & nutrition
– Induction of plant galls: abnormal structures
where gall makers (aphids, wasp, mites, etc.)
feed inside
• Manipulate plants’ defenses
Manipulate hostplants: induction of
plant galls
Copyright © 2007 Ilona Loser
Sumac galls
Aphids in sumac galls
Manipulate hostplants : repress toxinproduction
GOX=glucose oxidase
Musser et al., 2002, Nature
Plant-insect signaling interactions
• Are all plant defenses (or insect
counterdefenses) expressed (or present)
all the time? Why or why not?
• If yes, are they expressed at their highest
level all the time? Why or why not?
Plant-insect signaling interactions:
“spy” on each other and induction
Plant-insect signaling interactions: constitutive
vs. induced defense or counterdefense
Constitutive
l Present round-the-clock
l Predictable interactive
partners (e.g., specialist
herbivores)
l Pre-pay and fix the cost of
defense / counterdefense
no matter they are needed
or not
l Permanent protection. No
signal input required in
either side.
Induced
l Present or up-regulated only when
the interacting individuals
encounter each other
l Unpredictable interactive partners
(generalist herbivores)
l Defer the cost of
defense/counterdefense until they
are needed. But there is a lag time.
l Temporary protection. Signal input
from the other side needed
Plant-insect signaling interactions:
signals that plants receive from
herbivores and other plants
• Wounding: feeding or mechanical damages
• Herbivore-associated molecular patterns
(HAMPs)
– Chemicals: Fatty acid-amino acid conjugate
(FAC), caeliferins, 2-hydroxyoctadecatrienoic
acid, bruchins, and benzyl cyanide
– Enzymes: Glucose oxidase, β-glucosidase
– Proteolytic fragments of plant protein or
enzymes: e.g. inceptin (ICDINGVCVDA) from
plant chloroplastic
ATP synthase
• Talking tree: volatiles emitted from infested plants
Wounding and HAMPs trigger production of
endogenous plant defense signals
• Plant peptide hormones
– Systemin: mainly in the Solanaceae family
– HypSys pepetides: mainly in the Solanaceae family
– AtPep1: throughout the plant kingdom
• Plant defense signaling hormones
– JA (jasmonic acid), Ethylene (ET), SA (salicylic acid) :
well-characterized
– abscisic acid (ABA), auxin, gibberellic acid (GA),
cytokinin (CK), and brassinosteroids (BR)
Signals that plants received from
herbivores: wounding
O
COOH
•Wounding often leads to accumulation of plant defense signal molecules
such as JA, salicylic acid (SA), and ethylene (E), which are plant
hormones
• Plant defense hormones are systemic signals
•JA lead to production of the end defense products such as allelochemicals
and proteinase inhibitors (PIs)
JA: Jasmonic acid
Signals that plants received from herbivores: HAMP
JAburst Ethylene burst
Direct defense: nicotin
Kessler & Baldwin 2002
Elicitors modulate
defense response
Signals that plants received: volatiles
emitted from infested plants
• Dolch & Tscharntke (2000)
Field experiment:
Alnus
leaf
damage
(%)
eggs
per
leaf
distance
distance from defoliated tree (m)
• Volatile organic compounds (VOCs): terpenes, green leafy volatiles
• Methyljasmonate (MeJA)
• Talking tree
Plant-insect signaling interactions:
signals that insects received from plants
• Plant defense compounds (end products)
– Allelochemicals
– Proteinase inhibitors (PIs)
• Plant defense signaling hormones
– JA
– SA
– Ethylene
Signals insects received from plants :
allelochemicals and protease inhibitors
Li et al., 2002
Signals insects received from plants:
plant defense signaling hormone
0 30 min-5 hr
0 24hr
4 -6
days
Li et al., 2002, Nature
Types of plan-insect interactions
• Mutalism (+,+): flower plants/pollinators,
plants/ants
• Antagonistic herbivory (+,-): Insects eat
plants and plants suffer tissue lose, low
survival and reproduction.
• Antagonistic insectovory (+,-): plants eat
insects for nitrogen nutrients.
• Commensalism (+, 0):
Evolution of Resource Harvesting Organs with Resilience of Insects and Role of Plants for Sustenance of Insect Diversity

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Evolution of Resource Harvesting Organs with Resilience of Insects and Role of Plants for Sustenance of Insect Diversity

  • 1. EVOLUTION OF RESOURCE HARVESTING ORGANS WITH RESILIENCE OF INSECTS AND ROLE OF PLANTS FOR SUSTENANCE OF INSECT DIVERSITY AND MODES OF INSECT PLANT INTERACTION
  • 2. Introduction v Theinsecta are by far the most species-richtaxon. v Several hypotheses have been propounded to explain the evolution of the striking diversity of the Insecta and in particular of herbivorous insects. vAnew insect taxon could become established only if another was excluded by competition and becameextinct.
  • 3. Evolutionary diversity of insect ears •Insects are among the oldest land animals, and exist for more than 400 millionyears. •Among insects, tympanal earsevolved at least 18 times, resulting in adiversity of auditory systems. •Insects use their ears in different behavioural contexts, mainly intraspecific communication for mate attraction, predator avoidance, and parasitic host localisation.
  • 4. Evolutionary origin of auditory behaviour v It is driven by natural and sexual selection, insect ears evolved mainly for intraspecific communication and predator detection. v The most prominent example is between bats and nocturnal flying insects. v Ears of bats should be highly sensitive to ultrasound. Hemiptera of small body size use ancestrally tymbal mechanisms to generate vibration signals and Cicadidae produce auditory sounds by identical means of tymbal mechanisms.
  • 5. Ten regions where tympanal organs have evolved within insect groups 1. Lepidoptera; Sphingidae 2. Orthoptera; Ensifera 3. Diptera; Tachinidae 4. Mantodea; Mantidae 5. Lepidoptera; Geometroidea and Pyraloidea 6. Orthoptera; Acrididae 7. Hemiptera; Cicadidae 8. Lepidoptera; Noctuoidea 9. Hemiptera; Corixidae 10. Neuroptera; Chrysopidae
  • 6. Evolutionary origin of insect wings Theories of wing origin: 1.Paranotal theory 2.Epicoxal theory 3.Endite- Exite theory
  • 7.
  • 8.
  • 9.
  • 10.
  • 11.
  • 13.
  • 14. But lacking the tarsus, where the wing’s coastal surface normally would be.
  • 15. Role of Plants for sustenance of insect diversity Various factors have been proposed as agents of selection in the evolution of the specialistic feeding habit of herbivorous insects. Themain factors are: (1) coping with plant secondarymetabolites (2) avoiding competition (3) reducing mortality fromnatural enemies.
  • 16. Coping with plant secondary metabolites • Insects that are capable of detoxifying one class of plant compound usually cannot detoxify a very different class of secondary metabolites. • Specialist insects using the same plant taxon have evolved different detoxification or excretion mechanisms to avoid the impact of the samesecondary plantmetabolites. • These are steroidal compounds responsible for moulting, growth and maturation of insects. • It is produced by prothoracicgland. • An exciting investigation has shown that a single plant gene can determine whether the plant will be included in the diet of insect herbivores.
  • 17. Herbivory v Herbivory (phytophagy): Leafchewing , sapsacking, seedpredation, gall inducing, leafmining v Insect-plant mutualism –pollination and plantinsect food for defence relationshipsHerbivory v Chewersmost diverse of theleaf chewing insects are the Coleoptera and Lepidoptera. v Insects eat leaves,roots, shoots, stems, andflowers or fruits v Chewing insects possessmandibulate mouthparts v Mandibles serve to cut and grindfood v Mandibles are highly sclerotized to reducewear v High silica content and cellulose canact asresistance to herbivory
  • 18. Mining and boring • Insects live in between 2 epidermal layers of aleaf. Damageappears astunnels , blotches or blisters. • Independently evolved in 4 orders : Diptera,Lepidoptera, Coleoptera, and Hymenoptera • Different speciesmay excavate different layers of leaf parenchyma or reside in particularleaf • Fruit boring • Stem boring • Wood borers • Stalk boring • Plant boring
  • 20. Sap sucking • Drains plants resources by tapping into xylem and phloem • Hemipterans exemplify this strategy through haustellate mouth parts and Serves to pierce tissues and suckliquid food. • Labium modified into asheath enclosingstylet maxillae • Stylets pierce cuticle and canchangeorientation • Foodchannels empties into cibarialcavity
  • 22. Chewers • No relative sizerestrictions • Heavymechanical damage • Facedwith indigestible compounds andtoxins Suckers • Restricted to arelatively smallsize • Avoid mechanical damage (but still damaging) • Avoid indigestible compounds and mosttoxins • Xylem less suitable
  • 23.
  • 24. Gall makers • Gallsconsists of pathologically developed cells, tissues or organs of plants that have arisen by hypertrophy and/or hyperplasia asa result of stimulation from foreign organisms. • Orders that makesgalls;Hemiptera Diptera Hymenoptera
  • 25. Pollination • A pollinator moves pollen is an animal that from the male anther of aflower to the female stigma ofaflower. • This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollengrains. • Apollinator is different from apollenizer. • A plant that is a source of pollen for the pollination process.
  • 26. Types of pollinators Bees • Themost recognized pollinators are the various speciesof bees, which areplainly adapted to pollination. • Beestypically are fuzzy andcarry an electrostatic charge. • Both features help pollen grains adhere to their bodies, but theyalso have specialized pollen-carrying structures
  • 27. Honey bees • Honey bees travel from flower to flower, collecting nectar (later converted tohoney), and pollen grains. • The bee collects the pollen by rubbing against the anthers. • Nectar provides the energy for bee nutrition; pollen provides the protein. • Good pollination management seeks to have bees in a "building" state during the bloom period of the crop, thus requiring them to gather pollen, and making them more efficientpollinators.
  • 28. Other insects • Many insects other than beesaccomplish pollination by visiting flowers for nectar or pollen, or commonly both. • Many do so adventitiously, but the most important pollinators are specialists for at least parts of their lifecycles for at least certainfunctions. • Many bee flies, and someTabanidaeand Nemestrinidae are particularly adapted to pollinating fynbos and Karoo plants with narrow, deep corolla tubes, such as Lapeirousia species.
  • 29.
  • 30. Predation • Predation is a biological interaction where a predator feeds on its prey. • An insect predator is large in size, active in habits and have structural adoptions for catching the prey with well development senceorgans and capacity for swiftmovements. • Theimportant entomophagous predators are: Ø Dragonflies(Odonata): Thenaiads feed on aquatic insects and the adult on insects like mosquitoes, flies and moths while on the wings. Ø Praying mantids(Dictyoptera): feed upon flies, grasshopper and caterpillers. Ø Lady bird beetle(Coccinellidae): feed on soft body insects. Ø Tiger beetle( cicindellidae): Cicindella spp. feed upon avariety of insects. Ø Ground beetle(Carabidae): feed upon Opisina arenosella.
  • 32. Parasitism • Parasitism is anon-mutual relationship between specieswhere one species, the parasite, benefits at the expense of the other, the host. • Types:Insect parasitism is of fourkinds: a. Simple parasitism: Thisterm is applied when there is asingle attack of the parasitoid on the host.irrespective of the number of the eggs laid. b. Super parasitism: Parasitization of ahost by more than oneparasitic individual usually of one kind -used especially of parasitic insects. c. Multiple parasitism: Acondition in which parasites of differentspecies parasitize asingle host, in contrast to superparasitism or hyperparasitism. d. Hyperparasitism: It meansattack of aparasitiod on an insect which.is already aparasite asin the caseof bethylid and braconid parasites of Opisina arenosella.
  • 33. Mode of Insect Plant Interaction
  • 34. Overview • Brief introduction: herbivory & arms race • Arms / Weapons: – Diversity of plant defenses – Diversity of insect offence / counterdefense • Signaling interactions: – Signals that plants received from herbivores – Signals that insects perceived from plants
  • 35. Herbivory: host range Oliogophagy (<3 plant families ) Polyphagy Generalist Monophagy Specialist
  • 36. Herbivory: an arms race Insects attack consume plants Plants defend against being eaten warfare
  • 37. The arms race: coevolution Successful consumer New defenses Successful defender New offenses stepwise reciprocal changes
  • 38. The arms race: a typical example Insect resistant to toxin 3 Insect resistant to toxin2 Insect resistant to toxin1 Insect eats plant Plants make toxin 1 Plants make toxin 2 Plants make toxin 3 Plants make toxin 4
  • 39. Arms: diversity of plant defenses • Avoidance---escape from herbivores, no actual feeding. • Resistance---reduce fitness of insects after contact • Tolerance---to stand and take it simply by outgrowing the damage---compensatory growth
  • 40. Plant defenses: avoidance • Escape in time • Escape in space • Chemical escape (repellent production, no attractant) • Morphological escape • Heliconius butterflies avoid laying eggs on plants already occupied by eggs • Plants Passiflora create fake yellow eggs
  • 41. Plant defenses: Resistance • Morphological resistance – hairs, spines, hook, sticky glands, immobilizing insects or puncturing their body wall • Chemical resistance – Various toxic compounds • Mechanical resistance – squirt-gun
  • 42. Mechanical resistance: Bursera’s squirt-gun defense Becerra, J. X., 2003, PNAS 100 (22), 12804-12807
  • 43. Morphological resistance : thorns • The pronounced thorns endowed upon the trunk of this tree even can fend off vertebrate herbivores, such as sloths and monkeys.
  • 44. Chemical resistance: diversity & classification • Chronic (quantitative) defense: Large/complex molecules reduce digestibility / nutrition (tannins, lignins, cellulose, silica, etc). • Acute (qualitative) defense: Small / simple molecules target specific insect system. 1.Toxic amino acids 2.Toxic proteins 3. Proteinase inhibitors (=PIs) 4. Allelochemicals (=secondary comounds) cyanide, glycosides, alkaloids, terpenoids, saponins, flavenoids, furanocoumarins, indoles, phytoecdysteroids
  • 46. Arms: Insect’s offense/counterdefenses • Behavioral avoidance: selectively choose hostplants. feeding/oviposition choice • Modify its own behavior, or biochemistry to overcome plant defense • Behavioral counterdefense (or offense) • Biochemical counterdefense • Actively manipulate hostplants’s nutrition & defense
  • 47. Insect’s counterdefenses: Behavioral offense • Vein cutting • trenching • Leaf rolling • Mining • Gardening • Gregarious feeding
  • 48. Behavioral offense: vein cutting Specialized Blepharida’s counterdefense vein cutting Becerra, J. X., 2003, PNAS 100 (22), 12804-12807 Bursera’s defense squirt-gun
  • 49. Behavioral offense: trenching avoid intoxication by trenching the laticifers upstream of their intended feeding site (Wittstock & Gershenzon, 2002) Larvae of Erinnyis alope starting to feed after servering (trenching) a Carica papaya
  • 50. Behavioral offense: leaf-rolling Berenbaum & Zangerl: webworm-parsnip chemical co-evolution
  • 51. Behavioral offense: gardening l Leaf cutter ants (Azteca) use leaf material and flower to culture a fungal garden (facilitate food storage and toxin degradation), which is then used for food.
  • 52. Behavioral offense :gregarious feeding (group counterdefense) l Sunn Pest: gregarious feeding on wheat l increases host plant susceptibility l Pine beetle: calling for help if needed
  • 53. Insect’s counterdefenses: Biochemical counterdefense • Rapid excretion • Sequestration of toxins • Detoxification of toxins: cytochrome P450 monooxygenase (P450), esterase, glutathione-S-transferase (GST), etc. • Target site insensitivity
  • 54. Biochemical counterdefense: rapid excretion Experiment: Nicotine dose in artificial diet –> 93% excreted within 2 hours! Experiment: Nicotine injected into hemolymph Manduca sexta
  • 55. Biochemical counterdefense: sequestration • Unpalatable insects sequestrate allelochemicals for their own defense • Many also develop warning color or/and gregarious feeding
  • 57. Biochemical counterdefense: target site insensitivity Na+,K+-ATPase is sensitive to ouabain (a cardiac glycoside). In Monarch butterflies a single-point mutation resulted in insensitive ATPase. Other example: induction of proteases in herbivore gut, which are insensitive of proteinase inhibitors
  • 58. Insect’s counterdefenses: manipulate hostplants • Manipulate plants’ growth & nutrition – Induction of plant galls: abnormal structures where gall makers (aphids, wasp, mites, etc.) feed inside • Manipulate plants’ defenses
  • 59. Manipulate hostplants: induction of plant galls Copyright © 2007 Ilona Loser Sumac galls Aphids in sumac galls
  • 60. Manipulate hostplants : repress toxinproduction GOX=glucose oxidase Musser et al., 2002, Nature
  • 61. Plant-insect signaling interactions • Are all plant defenses (or insect counterdefenses) expressed (or present) all the time? Why or why not? • If yes, are they expressed at their highest level all the time? Why or why not?
  • 62. Plant-insect signaling interactions: “spy” on each other and induction
  • 63. Plant-insect signaling interactions: constitutive vs. induced defense or counterdefense Constitutive l Present round-the-clock l Predictable interactive partners (e.g., specialist herbivores) l Pre-pay and fix the cost of defense / counterdefense no matter they are needed or not l Permanent protection. No signal input required in either side. Induced l Present or up-regulated only when the interacting individuals encounter each other l Unpredictable interactive partners (generalist herbivores) l Defer the cost of defense/counterdefense until they are needed. But there is a lag time. l Temporary protection. Signal input from the other side needed
  • 64. Plant-insect signaling interactions: signals that plants receive from herbivores and other plants • Wounding: feeding or mechanical damages • Herbivore-associated molecular patterns (HAMPs) – Chemicals: Fatty acid-amino acid conjugate (FAC), caeliferins, 2-hydroxyoctadecatrienoic acid, bruchins, and benzyl cyanide – Enzymes: Glucose oxidase, β-glucosidase – Proteolytic fragments of plant protein or enzymes: e.g. inceptin (ICDINGVCVDA) from plant chloroplastic ATP synthase • Talking tree: volatiles emitted from infested plants
  • 65. Wounding and HAMPs trigger production of endogenous plant defense signals • Plant peptide hormones – Systemin: mainly in the Solanaceae family – HypSys pepetides: mainly in the Solanaceae family – AtPep1: throughout the plant kingdom • Plant defense signaling hormones – JA (jasmonic acid), Ethylene (ET), SA (salicylic acid) : well-characterized – abscisic acid (ABA), auxin, gibberellic acid (GA), cytokinin (CK), and brassinosteroids (BR)
  • 66. Signals that plants received from herbivores: wounding O COOH •Wounding often leads to accumulation of plant defense signal molecules such as JA, salicylic acid (SA), and ethylene (E), which are plant hormones • Plant defense hormones are systemic signals •JA lead to production of the end defense products such as allelochemicals and proteinase inhibitors (PIs) JA: Jasmonic acid
  • 67. Signals that plants received from herbivores: HAMP JAburst Ethylene burst Direct defense: nicotin Kessler & Baldwin 2002 Elicitors modulate defense response
  • 68. Signals that plants received: volatiles emitted from infested plants • Dolch & Tscharntke (2000) Field experiment: Alnus leaf damage (%) eggs per leaf distance distance from defoliated tree (m) • Volatile organic compounds (VOCs): terpenes, green leafy volatiles • Methyljasmonate (MeJA) • Talking tree
  • 69. Plant-insect signaling interactions: signals that insects received from plants • Plant defense compounds (end products) – Allelochemicals – Proteinase inhibitors (PIs) • Plant defense signaling hormones – JA – SA – Ethylene
  • 70. Signals insects received from plants : allelochemicals and protease inhibitors Li et al., 2002
  • 71. Signals insects received from plants: plant defense signaling hormone 0 30 min-5 hr 0 24hr 4 -6 days Li et al., 2002, Nature
  • 72. Types of plan-insect interactions • Mutalism (+,+): flower plants/pollinators, plants/ants • Antagonistic herbivory (+,-): Insects eat plants and plants suffer tissue lose, low survival and reproduction. • Antagonistic insectovory (+,-): plants eat insects for nitrogen nutrients. • Commensalism (+, 0):