History of C3 : C4 photosynthesis research

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After WWII at late 1940 at the University of California, Berkeley, the details of photosynthetic carbon metabolism were sorted out by the chemists Melvin Calvin, Andrew Benson, James Bassham and a score of students and researchers utilizing the carbon-14 isotope and paper chromatography techniques (Calvin 1989). The pathway of CO2 fixation by the algae Chlorella in a fraction of a second in light resulted in a 3 carbon molecule called phosphoglyceric acid (PGA). For that original and ground-breaking work, a Nobel Prize in Chemistry was awarded to Melvin Calvin 1961. In parallel, plant physiologists studied leaf gas exchanges using the new method of infrared gas analysis and a leaf chamber where the net photosynthetic rates ranged from 10 to 13 u mole CO2/square metere.sec., with the conclusion that all terrestrial plants having the same photosynthetic capacities that were light saturated at less than 50% of sunlight (Verduin 1953, Verduin et al. 1959). These rates were determined in potted plants grown indoors under low light intensity.

Later in 1958-1963 at Cornell University, field grown maize was reported to have much greater leaf photosynthetic rates of 40 u mol CO2/square meter.sec and was not saturated at near full sunlight (Hesketh and Musgrave 1962; Hesketh and Moss 1963). This higher rate in maize was almost double those observed in other species such as wheat and soybean, indicating that large differences in photosynthesis exist among higher plants. At the University of Arizona, detailed gas exchange research on more than 15 species of monocot and dicot uncovered for the first time that differences in leaf anatomy are crucial factors in differentiating photosynthetic capacities among species (El-Sharkawy 1965; El-Sharkawy and Hesketh 1965). In tropical grasses, including maize, sorghum, sugarcane, Bermuda grass and in the dicot amaranthus, leaf photosynthetic rates were around 38−40 u mol CO2/square meter.sec., and the leaves have two types of green cells, i. e. outer layer of mesophyll cells surrounding a tightly packed cholorophyllous vascular bundle sheath cells. This type of anatomy was termed Kranz anatomy in the 19th century by the botanist Gottlieb Haberlandt while studying leaf anatomy of sugarcane (Haberlandt 1904). Plant species with the greatest photosynthetic rates and Kranz anatomy showed no apparent photorespiration, very low CO2 compensation point, high optimum temperature, high stomatal resistances and lower mesophyll resistances for gas diffusion and rates never saturated at full sun light (El-Sharkawy 1965). The research at Arizona was designated Citation Classic by the ISI 1986 (El-Sharkawy and Hesketh 1986). These species was later termed C4 plants as the first stable compound of CO2 fixation in light has 4 carbon as malate and aspartate (Karpilov 1960; Kortschak et al. 1965; Hatch and Slack 1966). Other species that lack Kranz anatomy were termed C3 type such as cotton and sunflower, as the first stable carbon compound is the 3-carbon PGA acid. At 1000 ppm CO2 in measuring air, both the C3 and C4 plants had similar leaf photosynthetic rates around 60 u mole CO2/square meter.sec. indicating the suppression of phototorespiration in C3 plants (El-Sharkawy and Hesketh 1965, 1986).


  • Calvin, M. 1989. Forty years of photosynthesis and related activities. Phptosynth. Res. 21:3-16.
  • El-Sharkawy, M. A. 1965.Factors Limiting Photosynthetic Rates of Different Plant Species. Ph.D. Dissertation, The University of Arizona, Tucson, USA.
  • El-Sharkawy, M. A., and Hesketh, J. D. 1965. Photosynthesis among species in relation to characteristics of leaf anatomy and CO2 diffusion resistances. Crop Sci. 5:517-521.
  • El-Sharkawy, M. A., and Hesketh, J. D. 1986.Citation Classic-Photosynthesis among species in relation to characteristics of leaf anatomy and CO2 diffusion resistances. Curr. Cont./Agr.Biol.Environ.27:14-14. Online http://www.library.upenn/edu/classics1986/A1986C691300001.pdf.
  • Haberlandt, G. 1904.Physiologische Pflanzanatomie. Engelmann, Leipzig.
  • Hatch, M.D., and Slack, C. R. 1966.Photosynthesis by sugar-cane leaves. A new carboxylation reaction and the pathway of sugar formation. Biochem.J. 101:103-111.
  • Hesketh, J.D., and Moss, D. N. 1963.Variation in the response of photosynthesis to light. Crop Sci. 3:107-110.
  • Hesketh, J. D., and Musgrave, R.B. 1962. Photosynthesis under field conditions. IV. Light studies with individual corn leaves. Crop Sci. 2:311-315.
  • Karpilov, Y.S. 1960. The distribution of radioactvity in carbon-14 among the products of photosynthesis in maize. Proc. Kazan Agric. Inst. 14:15-24.
  • Kortschak, H.P., Hart, C.E., and Burr, G.O. 1965. Carbon dioxide fixation in sugarcane leaves. Plant Physiol. 40:209-213.
  • Verduin, J. 1953. A table of photosynthesis rates under optimal, near natural conditions. Am.J. Bot. 40:675-679.
  • Verduin, J., Whitwer, E. E., and Cowell, B.C. 1959. Maximal photosynthetic rates in nature, Science 130:268-269.