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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.

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Reduction 3. The PCR cycle can be divided into three primary stages : 1.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. Carboxylation 2. Regeneration . It is also referred to as the Calvin cycle.

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

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

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

(R + PR) True or gross photosynthesis (GP) is thus calculated by adding amount of mitochondrial = (respired CO2 + photorespired CO2) GP = AP + R + PR .The measured CO2 uptake in the light is termed apparent or net photosynthesis (AP) AP = GP .

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

Carbon assimilation pathway (C3.CAM) 2. Nutrient supply 6. Leaf area index 5. CO2 3. light 2. Pathological conditions 7.C4. Environmental factors 1. Leaf orientation 5. Temperature 4. Pollutants . Leaf age 3. Leaf angle 6. Morphology 4. Soil water Genetic factors 1.PRODUCTIVITY IS INFLUENCED BY A VARIETY OF GENETIC AND ENVIRONMENTAL FACTORS.

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 rate of photosynthesis continues to increase until it reaches light saturation (LS).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). As fluence rate increases. At fluence rates above the compensation point. The light compensation point for most plants falls somewhere in the range of 10 to 40 Qmol 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. .

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

which means that availability of CO2 is often a limiting factor in photosynthesis. regardless of fluence rate. 350 Ql/l is well below the CO2 saturation level for most C3 plants at normal fluence rates. both the maximum rate of photosynthesis and the CO2 saturation level increase. reduced photorespiration At higher fluence rates.035 percent by volume or 350 Ql/l. Increased substrate for the carboxylation reaction 2. increased photosynthetic rates with higher CO2 levels results from two factors: 1. most C4 plants appear to saturate at CO2 levels at or just above normal atmospheric concentrations. In C3 plants.The CO2 concentration of the atmosphere is relatively low and reasonably stable at about 0. Interestingly. . Competition with oxygen.

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. as much as by the intracellular CO2 concentration.The rate of photosynthesis is actually determined not by the ambient CO2 concentration. .

Here it is assumed the water supply is adequate and. 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. .Since CO2 diffusion rates depend in part on concentration gradients. consequently. stomatal CO2 conductance is not limiting.

hence. over that required to support carboxylation would represent an inefficient use of resources. which is in turn dependent on the electron transport reactions. but is saturated with respect to availability of the acceptor molecule RuBP However. .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. any excess generation of RuBP. the carboxylation capacity of the system.

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

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

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C4 plants enjoy some advantage over C3 plants with respect to photosynthesis and water stress because of their higher water use efficiency. Stomatal closure and the resultant decrease in CO2 supply due to water stress imposes a major limitation on 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. .SOIL WATER POTENTIAL The rate of photosynthesis declines under conditions of water stress. When this occurs in the presence of light for prolonged periods of time. this lack of CO2 supply may lead to photoinhibition of photosynthesis. and in cases of severe water stress may cease completely.

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

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

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

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5 v 109 t yr 1) of that total is accounted for by terrestrial ecosystems. Over the major portion of the oceans.PRIMARY PRODUCTIVITY ON A GLOBAL SCALE Total global productivity is estimated to be approximately 172 billion metric tons of dry organic matter per year. The remaining 32 percent is accounted for by marine ecosystems. land-based production is about five times greater than the oceans. Agriculture accounts for only 5. On an area basis. on the other hand. This difference is at least partly explained by differences in nutrient supply. retain a much larger nutrient capital in the soil and litter where they can continue to support growth and productivity. dead organisms and sinking particles carry nutrients out of the lighted zone near the surface where photosynthesis occurs. Productivity on land is more than twice that of the oceans on an area less than half as large. Approximately 68 percent (117. Terrestrial ecosystems.3 percent of global productivity. .

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

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