Dr. Stephen Malcolm, Department of Biological Sciences
– Lecture summary:
• The world is green - why?
• Explanatory theories:
– Classical plant defense theory :
» Dethier (1954); Fraenkel (1959); Ehrlich & Raven (1964).
– Apparency theory :
» Feeny (1976).
– Optimal defense theory :
» Rhoades & Cates (1976).
– Feeding specialization hypothesis :
» Futuyma (1976).
BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 4: Plant defense theory 1: history Slide - 1
(Figure 4-2).
– “ Somehow, millions of herbs, shrubs, and trees outgrow, outlast, or outdefend their herbivore enemies. One of the principal challenges of modern ecology is to understand how plants escape from herbivores in time and space.
”
– “ Plant defense theories provide the organizing principles.
”
BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 4: Plant defense theory 1: history Slide - 2
• “ The ease with which plants are located by herbivores provides one theoretical perspective ”
– Apparency theory .
• “ the ease with which plants mobilize resources to defend themselves provides another ”
– Resource availability theory .
• “ Both are grounded in the idea that plants evolve defensive capabilities, that are countered by adaptive evolution of animals that eat them ”
– Classical theory .
BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 4: Plant defense theory 1: history Slide - 3
• Dethier (1954) and Fraenkel (1959):
– Different insect species react differently to different plants with different secondary chemistry.
• Dethier and then Ehrlich & Raven (1964) first developed the “ classical theory of biochemical coevolution ” between plants and insects:
– In adapting to evolved plant defenses, insect herbivores lose some ability to detoxify allelochemicals of unrelated plants and so their diet breadth is narrowed.
BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 4: Plant defense theory 1: history Slide - 4
• “ The classical theory predicts that related taxa of insects become locked into a chemical arms race with related taxa of plants.
”
• “ Why do animals eat some plants, and not others?
”
– E.g.
Berenbaum's coumarins.
• Tables 7.1
& 7.2, Figs. 7.1
& 7.2.
– From Strong, Lawton & Southwood. 1984. Insects on Plants.
Community Patterns and Mechanisms.
Oxford: Blackwell.
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– Plants that are easily found ( “ apparent ” ) by herbivores evolve different kinds of chemical defenses from those that are difficult for animals to locate ( “ unapparent ” ).
– The distribution of qualitative ( toxic ) defenses and quantitative ( digestibility reducing ) defenses varies according to plant
“ apparency ” (Table 4-1, summary).
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7. Apparency Theory and temporal variation:
• Timing and synchrony are important (Fig. 4-3):
– So life history timing could also be an effective defense:
• E.g.
the November timing of oviposition by female winter moths and flight by males.
– Because leaves of apparent plants change in digestibility with time, e.g.:
• Increasing tannins and decreasing protein (Fig. 4-4).
• Decrease in grass digestibility with age (Fig. 4-7).
• Decrease in cherry and sorghum leaf nitrogen with age (Fig. 4.8) - grasses are apparent to grazers!
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• Specialists ( Pieris rapae ) on crucifers ( Brassica ) are much more affected by nutritional content
(nitrogen) than defensive chemical content
(glucosinolates such as sinigrin) - Fig. 4-5.
• But these plants avoid specialists by being hard to find and avoid generalists by having toxic chemical defenses.
• Is this logically inconsistent?
– cabbage to oak?
– aphid to elephant?
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– Patterns of chemical defense distribution (after
Futuyma, 1976) may be explained partially by variations in plant modularity (Table 4-2):
• More ephemeral tissues may have defenses characteristic of unapparent plants and less ephemeral tissues may have defenses like those in apparent plants.
• Alternatively, tissue age may be a better predictor of defense type.
BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 4: Plant defense theory 1: history Slide - 9
• Apparency theory partially held up for a test using persistent and pioneer tree species on Barro
Colorado Island in Panama:
– Tables 4-3 and 4-4 after Coley (1983).
• Mature leaves of persistents were grazed less
(0.04%/day) than those of pioneers (0.24%/day).
• Exception:
– Tannins were more prevalent in young trees for both pioneers and persistents.
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11. Chemical response by plants to herbivory:
– Wound-induced defenses in birch trees ( Betula spp.) reduced fitness of the geometrid moth
Epirrita autumnata perhaps through increased phenolic content (Fig. 4-9).
– There is some evidence that plants can compensate for herbivory (see Fig. 8.3
of
Hendrix data from Begon et al.
1990)
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• It is important to bear in mind the relevant target or organism that a plant must aim at to defend itself.
– For example, in the leaf cutter example the plant may have to target both the ant and its symbiotic fungus.
• Thus plants can produce effective fungicides to kill the "microbial farm" of the ants.
• But phenolics are useless against the ants because the fungus detoxifies them.
• Thus receptor sites are almost certainly crucial to understanding plant herbivore interactions.
• What do the plants target and why?
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– More butterfly species on unapparent herbs, but more moth species on apparent plants:
• Table 4-5 and Tables 1, 2 & 3 from Futuyma
(1976).
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14. Test of the Feeding Specialization Hypothesis:
– P. troilus :
• spicebush swallowtail eats only spicebush ( Lindera ) and sassafras ( Sassafras ).
– P. glaucus :
• tiger swallowtail eats 20 food plant species.
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• Spicebush vs tiger swallowtails:
– Eat 163 vs 54 mg/day/g of Lindera leaves
– At an efficiency of 21% vs 12%,
– To produce a relative growth rate of 33 vs 6 mg/day/g.
• Within a species, patterns of feeding specialization can be important and may lead to evolution of new species (Fig. 4-12).
• P. glaucus canadensis is probably a different species.
• Thus regional and local host specialization may be common.
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Figure 4-2: Effects of leaf removal on seed production after
1 year (left) and 2 years (right) in Piper arieianum shrubs.
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Table 7.1:
From, Strong,
Lawton &
Southwood,
1984.
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Table 7.2: From, Strong, Lawton &
Southwood, 1984.
BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 4: Plant defense theory 1: history Slide - 18
Number of plant species per genus in genera of Apiaceae with different chemistries.
More spp. in genera with angular and linear furanocoumarins
(Strong, Lawton &
Southwood, 1984: after Berenbaum,
1983).
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Figure 7.2:
Number of
Lepidopteran spp. associated with
Apiaceae that vary chemically
(Strong, Lawton &
Southwood, 1984: after
Berenbaum, 1983).
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BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 4: Plant defense theory 1: history Slide - 21
Figure 4-3:
Effects of oak bud burst timing (broken line) on synchrony with winter moth egg hatch and consequent larval mortality (stippling).
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Figure 4-4: Winter moth larval
size influenced by leaf age.
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Figure 4-7: Grass growth curve and changing
proportion digestible to herbivores.
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Figure 4-8: Seasonal changes in nutrient and cyanide content in black cherry and sorghum.
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Increase in larval feeding efficiency of the butterfly Pieris rapae with increased dietary nitrogen.
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Figure 4-9: Reduced growth survival and fecundity of larvae of the moth Epirrita autumnata fed leaves of birch trees that were damaged or had larval frass added.
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Figure 8.3: Compensation via reduced death rate in control and damaged flowers of wild parsnip, Pastinaca sativa (Begon, Harper & Townsend, 1990, 2nd ed.).
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From, Futuyma (1976)
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Table 2: Futuyma (1976)
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Figure 4-12: Geographical ranges of two races of
the tiger swallowtail, Papilio glaucus.
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