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The Consumer Connection
What does Ecological Stoichiometry have to
  say about consumption in food webs?




                Matt Whalen
              23 October 2012
Nutrients are good (and often limiting), so animals should seek them out.

While it was known that consumers can regulate homostasis with
respect to body composition of C, N, P, etc.,
         a cost of such regulation (via excretion) was not assumed

                                           Boersma & Elser 2006

                                           - Unimodal response of consumer
                                           growth rates to P levels among
                                           diverse taxa (5 arthropods, 1 snail,
                                           1 fish)

                                           - low-P animals may be more
                                           sensitive to excess P intake

                                                     P*Herb   P*Carnivore

                                            Growth



                                                          P in diet
So What?

   - could explain why some generalist herbivores consume large
   amounts of low-quality (low P) instead of more nutritious parts

   - limits to growth, additional stoichiometric constraint...what effect
   would this have on ecosystem models?

   - Why herbivores expected to be more sensitive? Adapted to
   acquiring and storing limiting nutrients, but perhaps not to deal with
   excess

   - We do know that excretion have energetic costs (think kidneys
   working against osmotic pressure), so why was this ignored?

   - form of recycled nutrients (organic vs. inorganic) and impact on
   autotrophic vs. heterotrophic processes?

   - implications for coexistence with additional nonlinearity of growth
   response to food quality?
Although homeostasis, fitness AND body composition can be affected by
food quality (locust example)




                                  Decreases in food quality could impair
                                  performace of herbivores, what would this
                                  mean for higher trophic levels?

                                  Low quality plant  low quality herbivores
                                   bad for carnivores?

                                  If eutrophication, how will bottom-up vs. top-
                                  down respond?

                                                     Raubenheimer & Simpson 2004 Ecology
Generalists may be able to respond to changes in food quality by changing
intake of food OR selectively grazing (either different species, or finding
tastier member of the same species)




                                             Here, Daphnia increase feeding
                                             (beat) rate for low quality foods,
                                             but decease their food intake with
                                             ever increasing food quality




                                                             Plath & Boersma 2001 Ecology
Balancing Diets – animals may selectively feed on food of different quality to
 reach their fitness optima, or at least compromise when optimum cannot be
 reached



Generalist and specialist locusts
reached similar target composition
by modifying intake in food mixtures

But, the patterns among food
monocultures (dashed lines) differed




                                                         Raubenheimer & Simpson 2004 Ecology
The evolutionary side:

How are animals adapted to deal with food quality in their environments?

- Gut length and architecture

- Extraoral digestion

- Generalism vs. specialism

- Feeding at multiple trophic levels

- Seasonal issues in dominance/succession of food groups. Population dynamics
of consumers, resting stages, etc.

- others?
The evolutionary side:


     Denno and Fagan argue that many predators (herbivore consumers) are
     N-limited and therefore would benefit from supplementing diet with N-rich
     predators (intraguild predation or cannabalism)

     They furthre argue that this possible benefit helps to explain the evolution
     of omnivory in the sense of carnivores eating herbivores and other
     carnivores
                                                                 Denno & Fagan 2003 Ecology



     Putting the cart before the horse? What about mixing of plants and
     animals?

     Herbivores are N-rich compared with plants, so consuming them seems
     like it would be selectively favored when plant resources are not
     unlimited, assuming you have the morphological/physiological capability

     Alternatively, if N or P is eaten in excess, might be easier to eat more
     lower quality food to balance C deficit
2 Cool Stories

1. Lupines and Lepidopteran herbivores




                         abundance
                          Herbivore
                                      Lupine density




                                      P:C                 P:C


With succession, increased plant densities leads to greater competition and lower
P content in plants. Translates to decreased herbivore fitness! (predation my also
be important here)
                                                                  Apple et al. 2009 PLoS ONE
2 Cool Stories

2. Spiders and Grasshoppers

A top-down perspective

-Spiders and their pred. cues
change nutrient constraints in
grasshoppers

- “Scared” hoppers have
higher metabolic rate

- increased demand for
digestible C to fuel energy
demand (but lots of crap C)

↑ C:N in hopper bodies
↑ C:N in fecal material
↑ C:N of plant community!

                                 Hawlena & Schmitz 2010 PNAS
Nutrient contents of food appear to be very important to consumers, and
can drive ecological interactions, patters of species abundance, evolution of
feeding strategies, etc.


So, what is the correct scale?

         - numbers of individuals
         - macromolecules (protein, carbs, lipids, etc.)
         - elements (N, P, C)

In what situations would each be favored?

Does the scale of the question (high-level ecosystem phenomena vs.
predator-prey interactions)?

Is this a strict decision, or can we combine scales in models when desired?

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Ecological Stoichiometry: The Consumer Connection: Matt Whalen

  • 1. The Consumer Connection What does Ecological Stoichiometry have to say about consumption in food webs? Matt Whalen 23 October 2012
  • 2. Nutrients are good (and often limiting), so animals should seek them out. While it was known that consumers can regulate homostasis with respect to body composition of C, N, P, etc., a cost of such regulation (via excretion) was not assumed Boersma & Elser 2006 - Unimodal response of consumer growth rates to P levels among diverse taxa (5 arthropods, 1 snail, 1 fish) - low-P animals may be more sensitive to excess P intake P*Herb P*Carnivore Growth P in diet
  • 3. So What? - could explain why some generalist herbivores consume large amounts of low-quality (low P) instead of more nutritious parts - limits to growth, additional stoichiometric constraint...what effect would this have on ecosystem models? - Why herbivores expected to be more sensitive? Adapted to acquiring and storing limiting nutrients, but perhaps not to deal with excess - We do know that excretion have energetic costs (think kidneys working against osmotic pressure), so why was this ignored? - form of recycled nutrients (organic vs. inorganic) and impact on autotrophic vs. heterotrophic processes? - implications for coexistence with additional nonlinearity of growth response to food quality?
  • 4. Although homeostasis, fitness AND body composition can be affected by food quality (locust example) Decreases in food quality could impair performace of herbivores, what would this mean for higher trophic levels? Low quality plant  low quality herbivores  bad for carnivores? If eutrophication, how will bottom-up vs. top- down respond? Raubenheimer & Simpson 2004 Ecology
  • 5. Generalists may be able to respond to changes in food quality by changing intake of food OR selectively grazing (either different species, or finding tastier member of the same species) Here, Daphnia increase feeding (beat) rate for low quality foods, but decease their food intake with ever increasing food quality Plath & Boersma 2001 Ecology
  • 6. Balancing Diets – animals may selectively feed on food of different quality to reach their fitness optima, or at least compromise when optimum cannot be reached Generalist and specialist locusts reached similar target composition by modifying intake in food mixtures But, the patterns among food monocultures (dashed lines) differed Raubenheimer & Simpson 2004 Ecology
  • 7. The evolutionary side: How are animals adapted to deal with food quality in their environments? - Gut length and architecture - Extraoral digestion - Generalism vs. specialism - Feeding at multiple trophic levels - Seasonal issues in dominance/succession of food groups. Population dynamics of consumers, resting stages, etc. - others?
  • 8. The evolutionary side: Denno and Fagan argue that many predators (herbivore consumers) are N-limited and therefore would benefit from supplementing diet with N-rich predators (intraguild predation or cannabalism) They furthre argue that this possible benefit helps to explain the evolution of omnivory in the sense of carnivores eating herbivores and other carnivores Denno & Fagan 2003 Ecology Putting the cart before the horse? What about mixing of plants and animals? Herbivores are N-rich compared with plants, so consuming them seems like it would be selectively favored when plant resources are not unlimited, assuming you have the morphological/physiological capability Alternatively, if N or P is eaten in excess, might be easier to eat more lower quality food to balance C deficit
  • 9. 2 Cool Stories 1. Lupines and Lepidopteran herbivores abundance Herbivore Lupine density P:C P:C With succession, increased plant densities leads to greater competition and lower P content in plants. Translates to decreased herbivore fitness! (predation my also be important here) Apple et al. 2009 PLoS ONE
  • 10. 2 Cool Stories 2. Spiders and Grasshoppers A top-down perspective -Spiders and their pred. cues change nutrient constraints in grasshoppers - “Scared” hoppers have higher metabolic rate - increased demand for digestible C to fuel energy demand (but lots of crap C) ↑ C:N in hopper bodies ↑ C:N in fecal material ↑ C:N of plant community! Hawlena & Schmitz 2010 PNAS
  • 11. Nutrient contents of food appear to be very important to consumers, and can drive ecological interactions, patters of species abundance, evolution of feeding strategies, etc. So, what is the correct scale? - numbers of individuals - macromolecules (protein, carbs, lipids, etc.) - elements (N, P, C) In what situations would each be favored? Does the scale of the question (high-level ecosystem phenomena vs. predator-prey interactions)? Is this a strict decision, or can we combine scales in models when desired?