Ecological Significance of the C4 Syndrome

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).

Resource: Introduction to Plant Physiology/ Williams G. Hopkins

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

C3
Photorespiration CO2 Compensation (1 CO2 1-1) Temperature optimum (oC) photosynthesis Rubisco PEPcase Quantum yield as a function of temperature Transpiration ratio Light saturation (mole photon m-2 s-1) Yes 20-100

C4
No 0-5

20-25 20-25
Declining 500 1000 400 500

30-45
30-35 Steady 200 350 Does not saturate

Resource: Introduction to Plant Physiology/ Williams G. Hopkins

CO2 Compensation
In a close environment, CO2 compensation concentration would be the stable CO2 concentration in air when CO2 uptake and evolution have come to equilibrium. For C3 plants, values fall into the range of 20 to 100 1 CO2 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

 However, although the activity of the glycolate pathway is very low in some C4 plants for example Sorghum bicolor (see picture) most C4 plants appear to have both peroxisomes and the metabolic machinery to support photorespiration. According to Bidwell (1983), The weight of evidence thus favors the conclusion that C4 plants do photo-respire, but at much reduced rates.
C4 leaves are an efficient CO2 absorber and effectively trap and recirculate any CO2 that might be produced in the leaves!!

Introduction to Plant Physiology/ Williams G. Hopkins

 The high level of CO2 developed in the bundle sheath cells would tend to suppress photorespiration by out-competing O2 for binding to Rubisco.

In addition, the anatomical and biochemical adaptation of C4 leaves ensure that any CO2 that might escape the bundle-sheath cell is trapped and re-assimilated by PEPcase in mesophyll cells, before it has the opportunity to escape from the leaves.
Introduction to Plant Physiology/ Williams G. Hopkins

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

 As a result, the quantum yield of photosynthesis in C3 plants tends to decline with 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/ Williams G. Hopkins

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 C3 Plants Triticum aestivum Encelia californica 0.055 0.067 0.050 0.059 0.041 0.046 25oC 35oC

C4 Plants Zea mays

Antriplex rosea

0.057
0.054

0.059
0.054

0.058
0.054

Introduction to Plant Physiology/ Williams G. Hopkins

Disadvantage of C4: Low-temperature sensitivity

While the are some cold-tolerance c4 species, most perform poorly if at all, at low temperature. Zea mays, for example, will not grow at temperature below 12-15oC. This lower limit for growth is probably set by the enzyme pyruvate, phosphate dikinase, which is cold labile and experiences a substantial loss of activity below 12oC
Introduction to Plant Physiology/ Williams G. Hopkins

In Availability of CO2 and Water

Photosynthesis in most situation is limited by availability of CO2 and water. In C3 plants, even moderate water stress will initiate closure of the stomata and reduce the available supply of CO2. The low CO2 compensation concentration of C4 plant means they can maintain higher rates of photosynthesis at lower CO2 level. Thus C4 plants gain an advantages over C3 plants when the stomata are partially closed to conserve during a period of water stress.
Introduction to Plant Physiology/ Williams G. Hopkins

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/ Williams G. Hopkins

 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 of photosynthesis while effectively conserving water.
Introduction to Plant Physiology/ Williams G. Hopkins

 Even under ideal condition, CO2 supply limits photosynthesis in C3 plants to the extent that light 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/ Williams G. Hopkins

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 less efficient.
Introduction to Plant Physiology/ Williams G. Hopkins

How can the lower photosynthetic efficiency of C4 plants be reconciled with their apparent higher activity?
Recall that C4 plants are native to tropical or subtropical habitats where there is usually an abundance of light. They can take advantage of some of this excess light to generate the ATP needed to run the C4 cycle, concentration of CO2, and increase net carbon assimilation.
Introduction to Plant Physiology/ Williams G. Hopkins

 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, high light and low water, the C4 syndrome confers a definite advantage.

Introduction to Plant Physiology/ Williams G. Hopkins

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 also fall 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

Terminologies
Terms 1. CO2 compensation concentration meaning Ambient CO2 concentration at which the rate of CO2 uptake (for photosynthesis) is balanced by the rate of CO2 evolution (by respiration) A short-lived continuation of a strong light dependent CO2 evolving process in the dark (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) 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

2. Post-illuminating CO2 burst (PIB)

3. Peroxisomes

Terms 4. Quantum yield

Meaning Quantum Yield can also be defined in fluorescence (emission) in which case the quantum yield is a measure of the efficiency with which absorbed light produces some effect

Introduction to Plant Physiology/ Williams G. Hopkins