Light, CO2 and the proper temperatures, the factors directly affects the photosynthesis in crop plants. Water and mineral elements also influence photosynthesis.
Factor # 1. Light:
Light response curves for leaves are illustrated in Figures 1.19 and 1.20. If there is no light, there is dark respiration, which for a leaf is usually 5 to 10% of CO2 uptake in bright light. As the light level gradually increases, photosynthesis increases to the light compensation level, which is the light level at which CO2 uptake is equal to CO2 evolution (the carbon exchange rate, or CER = 0).
If the light level continues to increase, there is less increase in CER for each unit increase in light level until the light saturation level is reached. Any increase in light level after this level will not significantly increase CER; therefore leaves are more efficient at utilizing light energy at low irradiance levels.
Species differ in their responses to light levels. Most C4 species (Fig. 1.20, curve A) are able to increase photosynthesis even at light levels equal to full sunlight, whereas most C3 species reach light saturation before full sunlight. Figure 1.20 illustrates that usually the lower the maximum CER, the lower the light level at which light saturation occurs.
It should be noted that even though C4 species often do not become light saturated and do use- high light levels better than C3 species, they use dimmer light more efficiently (CO2 uptake per unit of light) than bright light. For example, at 50 and 10% of full sun the CER is approximately 72 and 17%, respectively, of that at full sun; the most efficient use of light by CER is always at the lowest light levels. Efficiency is the slope of the light response curve.
Factor # 2. Carbon Dioxide:
Concentration in the Atmosphere:
Carbon dioxide is a gaseous component of air. Dry air contains 78% nitrogen (N2), 21% oxygen (O2), 0.93% argon (Ar), 0.034% (340 ppm) CO2, and traces of other gases. Although CO2 is at a low concentration, 85 to 92% of a plant’s dry weight is derived from CO2 uptake in photosynthesis.
Because of the burning of fossil fuels (which represent photosynthetic production millions of years ago) and the burning of forests, there has been an increasing CO2 concentration in the atmosphere. The projections for fossil fuel (primarily coal) utilization indicate an even larger increase in this concentration in the future.
Since CO2 causes a greenhouse effect by absorbing infrared bands of light, the higher concentration will cause the earth to retain more heat, which could increase mean global temperatures. Such an increase could influence the global weather pattern enough to change rainfall patterns and crop productive capabilities in many regions of the earth.
Under high-light conditions, most crop species show a linear response for leaf photosynthesis to CO2 levels above the current atmospheric concentration of 340 ppm. Crop yields could be increased considerably in an atmosphere enriched with CO2 up to 1500 ppm.
Although there is no current practical means of doing this under field conditions, CO2 enrichment has shown great benefits in greenhouses, not only increasing dry matter yield but also hastening plant development. It would be interesting to know how much the increase in CO2 concentration in the atmosphere in the past 100 years (from approximately 290 to 340 ppm) has increased crop yields and influenced crop maturation.
Leaf Resistances to CO2 Assimilation:
Carbon dioxide gets to the chloroplast by diffusion from the air through the stomata to the cell and then to the chloroplast.
Impediments to CO2 movement into and through the leaf do occur, and scientists have termed them resistances and quantified them.
rCO2 = ra + rs + rm …(1.1)
Where, rCO2 — CO2 exchange rate, ra = laminar resistance; rs — stomatal resistance; and rm = mesophyll resistance. Laminar resistance (ra) is the CO2 concentration at the leaf surface (also called the boundary layer effect), the lower the concentration, the higher and the resistance. Since ambient CO2 concentration is between 300 and 360 ppm, the factors that cause reduced concentration would increase ra.
In the field, turbulence is the major factor influencing ra. If no air movement occurs, CO2 uptake by the leaf causes a CO2 diffusion gradient that reduces CO2 concentration at the leaf surface. As wind turbulence increases, it will eventually reduce ra to the minimum level at most leaf surfaces within a plant canopy.
Stomatal resistance (rs) is the resistance of CO2 diffusion from outside the leaf through the stomata. Crop leaves generally have the stomatal frequency necessary for efficient CO2 diffusion. The major factor affecting rs is the degree to which stomata are open.
To calculate rs, crop physiologists measure the water loss from the leaf, which is a measure of stomatal impedance and diffusion. The rs is easily measured assuming that the relative humidity inside the leaf remains near saturation and any water loss is due to stomatal opening and ra.
Mesophyll resistance (rm) is calculated as residual resistance to CO2 uptake by the leaf:
rm = rCO2 – ra – rs
Mesophyll resistance is a measure of everything about the leaf that affects CO2 uptake except for ra and rs. This is because anything that influences CO2 fixation will affect CO2 concentration at the chloroplast, which in turn influences the total diffusion rate of CO2 from air to chloroplast.
The resistance formula (1.1) is used by crop physiologists as a method to determine if CO2 uptake by a crop plant is affected by resistance to CO2 diffusion into the leaf (ra and rs) or by CO2 fixation in the leaf (rm).
Factor # 3. Temperature:
Photosynthesis must be separated into its component parts to establish its response to temperature. The light reaction, or photophosphorylation, is independent of temperature in the temperature range in which plants grow. Carbon dioxide fixation is an enzymatically controlled reaction and increases at an increasing rate with increases in temperature until temperature reaches a level that favors enzymatic denaturization.
Respiration rates will continue to increase as temperature increases. Measurement of net CERs shows a minimal response of CER to temperature. Photorespiration also increases with temperature, since it is also enzymatically controlled, resulting in lower CER rates for C3 species than for C4 species at the higher temperatures of plant growth.
Factor # 4. Water:
Water is a substrate for photosynthesis, but only about 0.1% of the total water is used by the plant for photosynthesis. Transpiration accounts for 99% of the water used by plants; approximately 1% is used to hydrate the plant, maintain turgor pressure, and make growth possible. The primary influence of water stress on CER is an increase in r, due to stomatal closure. If water stress becomes severe, rm will also increase because of permanent damage to the photosynthetic apparatus.
Factor # 5. Leaf Age and Mineral Status:
Leaf age has an effect on photosynthesis- senescence causes a reduction in the process. The major factor that influences the rate of senescence is the mineral nutrient status of the leaf. Adequate mineral nutrient supplies allow both old and young leaves to meet their nutrient needs. However, limited nutrients are preferentially distributed to young leaves and reduce the photosynthetic rate of older leaves.
In maize, Peaslee and Moss (1966) measured lower photosynthetic rates for lower leaves. The lower rates were correlated with lower levels of potassium, phosphorus, magnesium, and nitrogen.
Apparently if these nutrients are in short supply they are translocated from older to younger leaves, causing what appears to be more rapid ageing in lower leaves. Other nutrients that are less mobile in the plant (e.g., calcium and iron) can reduce photosynthesis in younger leaves, while photosynthesis increases in older leaves due to the steady increase in calcium and iron content over time.
Reduced nutrient levels influence photosynthesis primarily by influencing the photosynthetic apparatus. For example, chlorophyll contains both nitrogen and magnesium; if they are limited in supply, chlorophyll may not form. Precursor molecules for chlorophyll synthesis include iron, and if it is not present, chlorophyll cannot be synthesized.
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