CARBON ASSIMILATION AND PRODUCTIVITY

Carbon assimilation creates plant biomass, or dry matter, which in turn supports humans and virtually all other heterotrophic organisms in the biosphere. The physiology of photosynthesis light capture, energy conversion, and carbon assimilation are at the root of productivity. PRODUCTIVITY REFERS TO AN INCREASE IN BIOMASS Although inorganic nutrients are a part of this dry matter, by far the bulk of dry matter for any organism consists of carbon.

PHOTOSYNTHETIC CARBON REDUCTION CYCLE (PCR)

The pathway by which all photosynthetic eukaryotic organisms ultimately incorporate CO2 into carbohydrate is known as carbon fixation or the photosynthetic carbon reduction (PCR) cycle. It is also referred to as the Calvin cycle. The PCR cycle can be divided into three primary stages : 1. Carboxylation 2. Reduction 3. Regeneration

THE CARBOXYLATION REACTION: How Calvin unraveled the path of carbon in photosynthesis?

The basic input into the biosphere is the conversion of solar energy into organic matter by photosynthetic plants and microorganisms, known as primary productivity (PP). Total carbon assimilation is known as gross primary productivity (GPP). Not all of the GPP is available for increased biomass - there Is a respiratory cost that must be taken into account. The principal focus of most productivity studies is therefore net primary productivity (NPP). NPP is determined by correcting GPP for energy and carbon loss due to respiration. NPP is a measure of the net increase in carbon, or carbon gain, and reflects the additional biomass that is available for harvest by animals.

CHLOROPLASTS OF C3 PLANTS ALSO EXHIBIT COMPETING CARBON OXIDATION PROCESS 1. The rate of photosynthesis is to measure gas exchange-either CO2 uptake or O2 evolution. 2. Cellular (or mitochondrial) respiration (R) is an opposite gas exchange, since it results in an evolution of CO2 and uptake of O2. 3. Evolution of CO2 associated with photosynthetic metabolism called photorespiration (PR) 4. PR involves the reoxidation of products assimilated in photosynthesis. 5. The photorespiratory pathway involves the activities of at least three different cellular organelles (the chloroplast, the peroxisome, and the mitochondrion) 6. CO2 evolved in PR results in a net loss of carbon from the cell.

The measured CO2 uptake in the light is termed apparent or net photosynthesis (AP) AP = GP - (R + PR)

True or gross photosynthesis (GP) is thus calculated by adding amount of mitochondrial = (respired CO2 + photorespired CO2) GP = AP + R + PR

CARBON ECONOMY
Carbon economy is the term used to describe the balance between carbon acquisition and its utilization. Respiration is the principal counterbalance to photosynthesis. Respiration consumes assimilated carbon in order to obtain the energy required to increase and maintain biomass. Respiratory loss of carbon constitutes one of the most significant intrinsic limitations on plant productivity.

PRODUCTIVITY IS INFLUENCED BY A VARIETY OF GENETIC AND ENVIRONMENTAL FACTORS.

Environmental factors 1. light 2. CO2 3. Temperature 4. Soil water Genetic factors 1. Carbon assimilation pathway (C3,C4,CAM) 2. Leaf age 3. Morphology 4. Leaf area index 5. Leaf angle 6. Leaf orientation 5. Nutrient supply 6. Pathological conditions 7. Pollutants

PHOTON FLUENCE RATE

Total number of photons (Np) incident from all directions on a small sphere divided by the cross-sectional area of the sphere and per time interval (m-2 s-1). At very low fluence rates the rate of CO2 evolution due to dark respiration exceeds the rate of photosynthetic CO2 uptake = negative CO2 uptake. As fluence rate increases, photosynthesis also increases and so does CO2 uptake until the rate of CO2 exchange equals zero = the light compensation point At the light compensation point the photosynthesis and the respiration are balanced. The light compensation point for most plants falls somewhere in the range of 10 to 40 Qmol m 2 s 1. At fluence rates above the compensation point, the rate of photosynthesis continues to increase until it reaches light saturation (LS).

In most C3 plants at normal atmospheric CO2 levels, photosynthesis saturates with light levels of about 500 to 1000 Qmol photons m 2 s 1, that is, about one-quarter to one-half of full sunlight. C4 plants never really achieve LS. There are also a small number of C3 plants, such as peanut (Arachis hypogea), which do not light saturate. The light saturated rate of photosynthesis, for example, is lower in leaves that have acclimated to growth at low irradiance (shade leaves) than in those that have acclimated to higher irradiance (sun leaves). Between dawn and dusk, the rate of photosynthesis gradually increases, reaching a maximum near midday, and then declines.

The CO2 concentration of the atmosphere is relatively low and reasonably stable at about 0.035 percent by volume or 350 Ql/l. 350 Ql/l is well below the CO2 saturation level for most C3 plants at normal fluence rates, which means that availability of CO2 is often a limiting factor in photosynthesis.

In C3 plants, increased photosynthetic rates with higher CO2 levels results from two factors: 1. Increased substrate for the carboxylation reaction 2. Competition with oxygen, reduced photorespiration At higher fluence rates, both the maximum rate of photosynthesis and the CO2 saturation level increase. Interestingly, most C4 plants appear to saturate at CO2 levels at or just above normal atmospheric concentrations, regardless of fluence rate.

The rate of photosynthesis is actually determined not by the ambient CO2 concentration, as much as by the intracellular CO2 concentration, that is, the supply of CO2 at the carboxylation site in the chloroplast. It assumed that the intracellular CO2 concentration is in equilibrium with the intercellular spaces.

Since CO2 diffusion rates depend in part on concentration gradients, the primary effect of increasing ambient CO2 levels would be to increase the intercellular CO2 concentration by increasing the rate of diffusion into the leaf. Here it is assumed the water supply is adequate and, consequently, stomatal CO2 conductance is not limiting.

Photosynthetic capacity is determined by the balance between carboxylation capacity and electron transport capacity

At low CO2 concentrations, the rate of photosynthesis is limited and, hence, the carboxylation capacity of the system, but is saturated with respect to availability of the acceptor molecule RuBP

However, any excess generation of RuBP, which is in turn dependent on the electron transport reactions, over that required to support carboxylation would represent an inefficient use of resources.

TEMPERATURE
Photosynthesis, like most other biological processes, is sensitive to temperature. The temperature response curve can be characterized by three cardinal points: the minimum and maximum temperatures (Tmin and Tmax, respectively) at which the reaction can proceed and the optimum temperature (Topt)
The temperature response of chemical and biological reactions can generally be characterized by comparing the rate of the reaction at two temperatures 10oC apart, a value known as the Q10: Q10 = RT +10 RT The value of Q10 for enzyme-catalyzed reactions is about 2, meaning that the rate of the reaction will approximately double for each 10oC rise in temperature. This value for Q10 applies primarily to stimulation of the reaction by temperatures between Tmin and Topt. Once the optimum is reached, the reaction rate may decline sharply due to enzyme inactivation

The temperature characteristics of C3 and C4 photosynthesis seem to be dominated by the temperature response curves for Rubisco and PEPcarboxylase, respectively. The low temperature sensitivity of C4 photosynthesis probably reflects, in part, the low temperature inactivation of the enzyme pyruvate, phosphate dikinase.

SOIL WATER POTENTIAL

The rate of photosynthesis declines under conditions of water stress, and in cases of severe water stress may cease completely. Stomatal closure and the resultant decrease in CO2 supply due to water stress imposes a major limitation on photosynthesis. When this occurs in the presence of light for prolonged periods of time, this lack of CO2 supply may lead to photoinhibition of photosynthesis. Photorespiration may protect the photosynthetic apparatus from excess light under such conditions because the energy absorbed can be used to fix O2 when the CO2 supply is limiting due to stomatal closure. C4 plants enjoy some advantage over C3 plants with respect to photosynthesis and water stress because of their higher water use efficiency.

NUTRIENT SUPPLY, PATHOLOGY, AND POLLUTANTS

Photosynthetic capacity is particularly sensitive to nitrogen supply. As a basic constituent of chlorophyll, redox carriers in the photosynthetic electron transport chain, and all of the enzymes involved in carbon metabolism, nitrogen plays a critical role in primary productivity. In a C3 species, Rubisco alone will account for more than half of the total leaf nitrogen. In one study of C3 and C4 grasses, net photosynthesis increased linearly with nitrogen content. In barley seedlings, a 5-fold increase in nitrate supply stimulated a 25-fold increase in net photosynthesis.

Photosynthesis energy is used directly in nitrate reduction and nitrogen assimilation provides amino acids for the synthesis of enzymes and proteins involved in both photosynthesis and respiration.

LEAF FACTORS
During initial development and the rapid growth phase, the photosynthetic capacity of leaf also increases. Leaf photosynthetic capacity then declines as the aging leaf undergoes senescence, a progressive deterioration of the leaf characterized in part by the loss of chlorophyll and photosynthetic enzymes. Different types of leaves may also have different photosynthetic capacities. Evergreen leaves, for example, have a lower photosynthetic capacity than deciduous leaves. The young, growing leaves at the top are exposed to full sunlight while older leaves further down may be heavily shaded. Very often the fluence rate reaching leaves lowermost in a canopy may fall below the light compensation point for a large part of the day.

The ratio of photosynthetic leaf area to covered ground area is known as the leaf area index (LAI). Because both leaf surface and the covered ground are measured as areas (m2), LAI is dimensionless. Values of LAI in productive agricultural ecosystems typically fall in the range of 3 to 5. The optimum LAI for a given stand of plants depends on the angle between the leaf and the stem. Horizontal leaves, typical of beans (Phaseolus) and similar crops, are efficient light absorbers because of the broad surface presented to the sun, but they also more effectively shade leaves lower down in the canopy. Erect leaves, typical of grasses like wheat (Triticum) and maize (Zea mays), produce less shading but, because of their steeper angle, are not as efficient at intercepting light. Field studies have confirmed that canopies with predominantly horizontal leaves have LAI values of 2 or less, while vertical leaf canopies support LAI values of 3 to 7.

PRIMARY PRODUCTIVITY ON A GLOBAL SCALE Total global productivity is estimated to be approximately 172 billion metric tons of dry organic matter per year. Approximately 68 percent (117.5 v 109 t yr 1) of that total is accounted for by terrestrial ecosystems. The remaining 32 percent is accounted for by marine ecosystems. Productivity on land is more than twice that of the oceans on an area less than half as large. On an area basis, land-based production is about five times greater than the oceans. This difference is at least partly explained by differences in nutrient supply. Over the major portion of the oceans, dead organisms and sinking particles carry nutrients out of the lighted zone near the surface where photosynthesis occurs. Terrestrial ecosystems, on the other hand, retain a much larger nutrient capital in the soil and litter where they can continue to support growth and productivity. Agriculture accounts for only 5.3 percent of global productivity.

SUMMARY
1. Carbon assimilation by plants creates the plant biomass that supports humans and virtually all other heterotrophic organisms. 2. Overall carbon gain depends on net primary productivity-the balance between carbon uptake by photosynthesis and carbon loss respiration. 3. Carbon loss to respiration, can be divided into the carbon cost of growth, or growth respiration, and the cost of simply maintaining structure and processes that do not result in a net increase in dry matter. 4. Several studies have shown a negative correlation between respiration and growth rate. 5. Productivity is also influenced by a variety of genetic and environmental factors that influence photosynthesis. These include light, available carbon dioxide, temperature, soil water, nutrients, and canopy structure.

6. Productivity is also influenced by a variety of genetic and environmental factors that influence photosynthesis. These include light, available carbon dioxide, temperature, soil water, nutrients, and canopy structure. 7. Plants also require an adequate water and nitrogen supply in order to maximize their leaf photosynthetic capacity. 8. Productivity depends on the pattern of leaf senescence and the structure of the canopy. The ideal canopy maximizes the efficiency of light interception and carbon gain by balancing leaf area, leaf angle, leaf orientation, plant density, and senescence of older leaves.