Lecture 3: Ecological Limiting Factors and Adaptation

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Reading:  Economy of Nature, pp. 60-70.

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Water absorbs light energy and scatters light

In sea water:   At 10m, the energy of visible light decreases 50%
                         At 100m, the energy of visible light decreases to <7%

Red is absorbed first
Blue and violet scatter easily
Green penetrates water best

Euphotic zone: Depth to which photosynthesis exceeds respiration in water.

Rarely the compensation point, the bottom of the euphotic
zone, is as deep as 100m. Examples, very clear ocean or
lakes near equator.
In highly turbid waters, the compensation point may be reached
at 1m.

Major Essential Elements

Calcium (Ca), Iron (Fe), Nitrogen (N), Magnesium (Mg), Potassium (K), Phosphorus (P), Sodium (Na), Sulfur (S)

Limiting Nutrient Elements

In aquatic (freshwater) environments: nitrogen and phosphorus
In marine (saltwater) environments: iron
In terrestrial environments: nitrogen and phosphorus (calcium)

Other Essential Resources

Carbon Dioxide: Not limiting

Oxygen: Can be limiting in water

Water: Often limiting in terrestrial environments

Limitations for one essential resources can influence the availability of other essential resources. This is the case among terrestrial plants for the relationships between photosynthetic rates, water loss, and gas exchange.

Photosynthetic Capacity and Water Conservation

Photosynthesis rate varies widely among species (100x) even with light saturation and all other resources in abundance. This variation is due in part to differences between plant species in the biochemistry of carbon fixation in photosynthesis (Calvin Cycle). Plants can be categorized as having , , or CAM metabolism.

Photosynthetic Rate, Water Loss and Gas Exchange Specializations and Compromises Among Plants

  • Short life, high photosynthetic rate when water abundant, dormant
    at other times (seed stage) (desert annuals)
  • Long life, leaves produced when water abundant, leaf drop during
    droughts (winter or dry season) (deciduous woody plants)
  • Leaves long lived, transpire slowly, tolerate water deficit but have
    low photosynthetic capacity (woody evergreens, evergreen desert
    shrubs)
  • photosynthesis: increased efficiency of carbon dioxide use per
    unit of water loss, but inefficient at low light intensity (not shade
    tolerant), high temperature optima, adaptation for water conservation
    and efficient nutrient capture (arid, tropical, and saline environments)
    (see figures on next page)
  • CAM photosynthesis (Crassulacean Acid Metabolism): control of
    water loss by limiting atmospheric carbon dioxide capture to night
    hours when water transpiration rates are at a minimum, stomata
    are open at night and are closed during the day, good water
    conservation but there are limits on photosynthetic capacity (arid,
    high elevation, windy environments)

Major steps in the capture of carbon dioxide in CAM plants. Atmospheric carbon dioxide is brought into the plant at night, when the stomata are open. In the daytime, stored reserves of malic acid and oxaloacetate are broken down to release carbon dioxide inside leaves for use in the Calvin Cycle, but gas exchange to the atmosphere is minimal. Abbreviation key: RuBP = ribulose bisphosphate (a 5C compound), PGA = phosphoglycerate (two molecules of a 3C compound), PEP = phosphoenolpyruvate (a 3C compound), OAA = oxaloacetate (a 4C compound), Malate = malic acid (a 4C compound). Water loss is controlled in this system by mean of temporal separation of carbon dioxide capture from the atmosphere and the addition of one carbon to RuBP in the first step of the Calvin Cycle (Ricklefs, 1996, p 69, Fig. 3.9).

 

 

Comparison between and plants showing the differences in the physical distribution of chloroplasts in the leaves, and differences in first steps of atmospheric carbon dioxide capture for photosynthesis. Abbreviation key: RuBP = ribulose bisphosphate (a 5C compound), PGA = phosphoglycerate (two molecules of a 3C compound), PEP = phosphoenolpyruvate (a 3C compound), OAA = oxaloacetate (a 4C compound), Pyr = pyruvate (a 3C compound). Water loss is better controlled in plants by physically separating carbon dioxide capture from the atmosphere and the addition of one carbon to RuBP in the first step of the Calvin Cycle. Water loss is minimized in this system because the atmospheric carbon dioxide capture step is catalyzed by the enzyme PEP carboxylase which has a higher affinity for carbon dioxide than does the enzyme RuBP carboxylase (Ricklefs, 1996, pp 68 and 69, Fig. 3.7 and 3.8).

Plants

Plants

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