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Ecological Significance

Apr 04, 2018

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Jauha Hanafi
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    Unlike C3 plants, photosynthesis of C4

    plants is not inhibited by O2 and they

    exhibit no post-illumination CO2 burst

    and have a very low CO2 Compensation

    concentration (Table 1).

    Ecological Significance of the C4

    Syndrome

    Resource: Introduction to Plant Physiology/ Williams G. Hopkins

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    C3 C4

    Photorespiration Yes No

    CO2 Compensation (1 CO2 1-1) 20-100 0-5

    Temperature optimum (oC)

    photosynthesis

    Rubisco

    PEPcase

    20-25

    20-25

    30-45

    30-35

    Quantum yield as a function of temperature Declining Steady

    Transpiration ratio 500 1000 200 350

    Light saturation (mole photon m-2 s-1) 400 500 Does not

    saturate

    Table 1: A comparison significant features of C3 and C4 plants

    Resource: Introduction to Plant Physiology/ Williams G. Hopkins

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    CO2 Compensation

    In a close environment, CO2 compensation concentrationwould be the stable CO2 concentration in air when CO2uptake and evolution have come to equilibrium.

    For C3 plants, values fall into the range of 20 to 100 1CO2 per liter.

    Comparable values for C4 plants are in range of 0-5 1-1

    In conclusion, photorespiration is either absent from C4

    plants or that the process is suppressed.

    Resource: Introduction to Plant Physiology/ Williams G. Hopkins

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    The high level of CO2 developed in the bundle

    sheath cells would tend to suppressphotorespiration by out-competing O2 forbinding to Rubisco.

    In addition, the anatomical and biochemicaladaptation of C4 leaves ensure that any CO2that might escape the bundle-sheath cell is

    trapped and re-assimilated by PEPcase inmesophyll cells, before it has the opportunityto escape from the leaves.

    Introduction to Plant Physiology/ Williams G. Hopkins

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    Temperature optimum

    Most C4 plants tend to have a higher

    temperature optimum (30-45oC) than C3

    plants (20-25oC)

    Due to higher temperature stability of some of

    the C4 cycle enzymes.

    Maximal activity of:

    PEPcase -> 30-35oC

    Rubisco -> 20-25oC

    Introduction to Plant Physiology/ Williams G. Hopkins

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    As a result, the quantum yield of

    photosynthesis in C3 plants tends to declinewith the increasing leaf temperatures due to

    decrease carboxylation activity of Rubisco at

    the higher temperatures

    While the quantum yield of C4 plants remains

    essentially constant.

    Introduction to Plant Physiology/ WilliamsG. Hopkins

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    Effect of temperature on quantum yield of

    photosynthesis in C3 and C4 plants(From Hopkins, W.G, Introduction of Plant Physiology. 203, 1995)

    Quantum yield (mol CO2 / mol quantana)

    15oC 25oC 35oC

    C3 Plants

    Triticum aestivum 0.055 0.050 0.041

    Encelia californica 0.067 0.059 0.046

    C4 Plants Zea mays 0.057 0.059 0.058

    Antriplex rosea 0.054 0.054 0.054

    Introduction to Plant Physiology/ WilliamsG. Hopkins

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    Disadvantage of C4:

    Low-temperature sensitivity

    While the are some cold-tolerance c4 species,most perform poorly if at all, at lowtemperature.

    Zea mays, for example, will not grow attemperature below 12-15oC.

    This lower limit for growth is probably set by

    the enzyme pyruvate, phosphate dikinase,which is cold labile and experiences asubstantial loss of activity below 12oC

    Introduction to Plant Physiology/ WilliamsG. Hopkins

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    Photosynthesis in most situation is limited byavailability ofCO2 and water.

    In C3 plants, even moderate water stress willinitiate closure of the stomata and reduce theavailable supply of CO2.

    The low CO2 compensation concentration of C4plant means they can maintain higher rates ofphotosynthesis at lower CO2 level.

    Thus C4 plants gain an advantages over C3 plantswhen the stomata are partially closed to conserveduring a period of water stress.

    Introduction to Plant Physiology/ WilliamsG. Hopkins

    In Availability of CO2 and Water

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    Transpiration Ratio

    The transpiration ratio (TR) relates to uptakes

    of CO2 to the loss of water by evaporation

    (transpiration) from the leaves.

    Introduction to Plant Physiology/ WilliamsG. Hopkins

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    Transpiration ratios for C4 plants are typically

    in the range of 200 350

    While for C3 plants values the range of 500

    1000

    The low transpiration ratio for C4 plants

    reflect their capacity to maintain high rates ofphotosynthesis while effectively conserving

    water.

    Introduction to Plant Physiology/ WilliamsG. Hopkins

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    Even under ideal condition, CO2 supply limits

    photosynthesis in C3 plants to the extent thatlight saturation occurs at fluence rate about

    25% of sull sunlight.

    C4 plants, on the other hand never really

    saturate, even at full sunlight.

    Introduction to Plant Physiology/ WilliamsG. Hopkins

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    Disadvantage of Respiration ration of

    C4 Plants

    C4 is not necessarily more efficient than C3

    photosynthesis

    At leaf temperature below 30oC, the quantum

    yield for C4 plants is actually lower than C3

    plant that is C4 photosynthesis is lessefficient.

    Introduction to Plant Physiology/ WilliamsG. Hopkins

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    How can the lower photosynthetic

    efficiency of C4 plants be reconciled with

    their apparent higher activity?

    Recall that C4 plants are native to tropical orsubtropical habitats where there is usually anabundance of light.

    They can take advantage of some of this excess

    light to generate the ATP needed to run the C4cycle, concentration of CO2, and increase netcarbon assimilation.

    Introduction to Plant Physiology/ WilliamsG. Hopkins

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    Although C4 plants are not competitive in all

    situations some C3 plant may even equal or

    exceed C4 plants in productivity given the

    right combination of high temperature, highlight and low water, the C4 syndrome confers

    a definite advantage.

    Introduction to Plant Physiology/ WilliamsG. Hopkins

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    Prove?

    The frustration of home owner and farmers as

    Many of our aggressive weeds are C4 species :

    crabgrass (Digitaria sanguinalis), Russian Thristle

    (Salsola kali), and several species of pigweed(Amaranthus) that often take over during the hot dry

    month in the middle of summer.

    Many of the more highly productive crop species alsofall within the C4 group, including sugarcane

    (Saccharum officinarum), sorghum (Sorghum bicolor,

    maize (Zea mays) and millet (Panicum miliaceum)Introduction to Plant Physiology/ Williams

    G. Hopkins

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    Terminologies

    Terms meaning

    1. CO2 compensation concentration Ambient CO2 concentration at which the

    rate of CO2 uptake (for photosynthesis) is

    balanced by the rate of CO2 evolution (by

    respiration)

    2. Post-illuminating CO2 burst (PIB) A short-lived continuation of a strong light

    dependent CO2 evolving process in thedark (e.g. Dekker 1975, Tregunna et al.

    1966)

    Later was taken as a measure of photo

    respiration rate and has been referred to

    glycolate pathway (e.g Chollet & Ogren

    1975)

    3. Peroxisomes A cell organelle containing enzymes, such

    as catalase and oxidase, that catalyze the

    production and breakdown of hydrogen

    peroxide.

    Introduction to Plant Physiology/ Williams G. Hopkins

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    Terms Meaning

    4. Quantum yield Quantum Yield can also be defined in

    fluorescence (emission) in which case the

    quantum yield is a measure of theefficiency with which absorbed light

    produces some effect

    Introduction to Plant Physiology/ WilliamsG. Hopkins