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Temperature x CO2 Interaction Plant Growth Response (Agricultural Crops)
As the airs CO2 content continues to rise, most plants will exhibit increased rates of photosynthesis and biomass production.  Consequently, this phenomenon should enhance the amount of food, fiber and timber products that can be used to feed, clothe and shelter earths expanding human population.  However, some individuals have predicted that the growth-promoting effects of atmospheric CO2 enrichment may be negated by global warming, which could compromise our ability to sustain greater human populations without increasing arable land acreage.  We thus turn to the scientific literature to see if plants will continue to exhibit CO2-induced growth increases under conditions of elevated air temperature.  In this summary, we review the photosynthetic and growth responses of agricultural crops within this context.

In order to better understand the issues being addressed in this review, one must realize that the optimum growth temperature for several plants has already been shown to rise substantially with increasing levels of atmospheric CO2 (McMurtrie and Wang, 1993; McMurtrie et al., 1992; Stuhlfauth and Fock, 1990; Berry and Bjorkman, 1980).  This phenomenon was predicted by Long (1991), who calculated from well-established plant physiological principles that most C3 plants should increase their optimum growth temperature by approximately 5C for a 300 ppm increase in the airs CO2 content.  One would thus also expect plant photosynthetic rates to rise with concomitant increases in the airs CO2 concentration and temperature, as has indeed been previously shown to be true by Idso and Idso (1994).  Hence, we here proceed to see if these positive CO2 x temperature interactions are still being supported in the recent scientific literature.

In the study of Zhu et al. (1999), pineapples grown at 700 ppm CO2 assimilated 15, 97 and 84% more total carbon than pineapples grown at the current ambient CO2 concentration in day/night air temperature regimes of 30/20 (which is optimal for pineapple growth at ambient CO2), 30/25, and 35/25 C, respectively.  Similarly, Taub
et al. (2000)
demonstrated that net photosynthetic rates of cucumbers grown at twice-ambient levels of atmospheric CO2 and air temperatures of 40C were 3.2 times greater than those displayed by control plants grown at ambient CO2 and this same elevated air temperature.  Thus, at air temperatures normally considered to be deleterious to plant growth, rates of photosynthesis are typically considerably greater for CO2 enriched vs. ambiently-grown plants.

Other studies report similar results.  Reddy et al. (1999), for example, grew cotton plants at air temperatures ranging from 2C below to 7C above ambient air temperatures and reported that plants simultaneously exposed to 720 ppm CO2 displayed photosynthetic rates that were 137 to 190% greater than those displayed by plants exposed to ambient CO2 concentrations across this temperature spectrum.  Similarly, Cowling and Sage (1998) reported that a 200-ppm increase in the airs CO2 concentration boosted photosynthetic rates of young bean plants by 58 and 73% at growth temperatures of 25 and 36C, respectively.  In addition, Bunce (1998) grew wheat and barley at 350 and 700 ppm CO2 across a wide range of temperatures and reported that elevated CO2 stimulated photosynthesis in these species by 63 and 74%, respectively, at an air temperature of 10C and by 115 and 125% at 30C.  Thus, the percentage increase in photosynthetic rate resulting from atmospheric CO2 enrichment often increases substantially with increasing air temperature.

On another note, elevated CO2 often aids in the recovery of plants from high temperature-induced reductions in photosynthetic capacity, as noted by Ferris et al. (1998), who grew soybeans for 52 days under normal air temperature and soil water conditions at atmospheric CO2 concentrations of 360 and 700 ppm, but then subjected them to an 8-day period of high temperature and water stress.  After normal air temperature and soil water conditions were restored, the CO2-enriched plants attained photosynthetic rates that were 72% of their unstressed controls, while stressed plants grown at ambient CO2 attained photosynthetic rates that were only 52% of their respective controls.

CO2-induced increases in plant growth under high air temperatures have also been observed in a number of other agricultural species.  In the previously mentioned study of Cowling and Sage (1998), for example, the 200-ppm increase in the airs CO2 content boosted total plant biomass for wheat and barley by a combined average of 59 and 200% at air temperatures of 25 and 36C.  Similarly, Ziska (1998) reported that a doubling of the atmospheric CO2 concentration increased the total dry weight of soybeans by 36 and 42% at root zone temperatures of 25 and 30C, respectively.  Likewise, Hakala (1998) noted that spring wheat grown at 700 ppm CO2 attained total biomass values that were 17 and 23% greater than those attained by ambiently-grown plants exposed to ambient and elevated (ambient plus 3C) air temperatures.  In addition, after inputting various observed CO2-induced growth responses of winter wheat into plant growth models, Alexandrov and Hoogenboom (2000) predicted 12 to 49% increases in wheat yield in Bulgaria even if air temperatures rise by as much as 4C.  Finally, in the study of Reddy et al. (1998), it was shown that elevated CO2 (700 ppm) increased total cotton biomass by 31 to 78% across an air temperature range from 20 to 40C.  Thus, the beneficial effects of elevated CO2 on agricultural crop yield is often enhanced due to elevated air temperature.

In some cases, however, elevated CO2 does not interact with air temperature to further increase the growth-promoting effects of atmospheric CO2 enrichment, but simply allows the maintenance of the status quo.  In the study of Demmers-Derks et al. (1998), for example, sugar beets grown at 700 ppm CO2 attained 25% more biomass than ambiently-grown plants, regardless of air temperature, which was increased by 3C.  Similarly, in the study of Fritschi et al. (1999), elevated CO2 concentrations did not significantly interact with air temperature (4.5C above ambient) to impact the growth of rhizoma peanut.  Nonetheless, the 300-ppm increase in the airs CO2 content increased total biomass by 52%, regardless of air temperature.

Finally, even if the airs CO2 content were to cease rising or have no effect on plants, it is possible that temperature increases alone would promote plant growth and development.  This was the case in the study of Wurr et al. (2000), where elevated CO2 had essentially no effect on the yield of French bean.  However, a 4C increase in air temperature increased yield by approximately 50%.

In conclusion, the recent scientific literature continues to indicate that as the airs CO2 content continues to rise, agricultural crops will likely exhibit enhanced rates of photosynthesis and biomass production that will not be diminished by any global warming that might occur concurrently.  In fact, if the ambient air temperature rises, the growth-promoting effects of atmospheric CO2 enrichment will likely rise right along with it, becoming more and more robust in agreement with the experimental observations reviewed by Idso and Idso (1994).  Thus, the biospheres ability to continue producing the food and fiber needed to feed and cloth the increasing population of humanity looks good indeed as long, that is, as the CO2 content of the air continues its 350-year habit of rising hand-in-hand with the population of the planet (see our Editorial of 29 August 2001).

Alexandrov, V.A. and Hoogenboom, G.  2000.  The impact of climate variability and change on crop yield in Bulgaria.  Agricultural and Forest Meteorology 104: 315-327.

Berry, J. and Bjorkman, O.  1980.  Photosynthetic response and adaptation to temperature in higher plants.  Annual Review of Plant Physiology 31: 491-543.

Bunce, J.A.  1998.  The temperature dependence of the stimulation of photosynthesis by elevated carbon dioxide in wheat and barley.  Journal of Experimental Botany 49: 1555-1561.

Cowling, S.A. and Sage, R.F.  1998.  Interactive effects of low atmospheric CO2 and elevated temperature on growth, photosynthesis and respiration in Phaseolus vulgarisPlant, Cell and Environment 21: 427-435.

Demmers-Derks, H., Mitchell, R.A.G., Mitchell, V.J. and Lawlor, D.W.  1998.  Response of sugar beet (Beta vulgaris L.) yield and biochemical composition to elevated CO2 and temperature at two nitrogen applications.  Plant, Cell and Environment 21: 829-836.

Ferris, R., Wheeler, T.R., Ellis, R.H. and Hadley, P.  1999.  Seed yield after environmental stress in soybean grown under elevated CO2Crop Science 39: 710-718.

Ferris, R., Wheeler, T.R., Hadley, P. and Ellis, R.H.  1998.  Recovery of photosynthesis after environmental stress in soybean grown under elevated CO2Crop Science 38: 948-955.

Fritschi, F.B., Boote, K.J., Sollenberger, L.E., Allen, Jr. L.H. and Sinclair, T.R.  1999.  Carbon dioxide and temperature effects on forage establishment: photosynthesis and biomass production.  Global Change Biology 5: 441-453.

Hakala, K.  1998.  Growth and yield potential of spring wheat in a simulated changed climate with increased CO2 and higher temperature.  European Journal of Agronomy 9: 41-52.

Idso, K.E. and Idso, S.B.  1994.  Plant responses to atmospheric CO2 enrichment in the face of environmental constraints: A review of the past 10 years research.  Agricultural and Forest Meteorology 69: 153-203.

Long, S.P.  1991.  Modification of the response of photosynthetic productivity to rising temperature by atmospheric CO2 concentrations: Has its importance been underestimated?  Plant, Cell and Environment 14: 729-739.

McMurtrie, R.E. and Wang, Y.-P.  1993.  Mathematical models of the photosynthetic response of tree stands to rising CO2 concentrations and temperatures.  Plant, Cell and Environment 16: 1-13.

McMurtrie, R.E., Comins, H.N., Kirschbaum, M.U.F. and Wang, Y.-P.  1992.  Modifying existing forest growth models to take account of effects of elevated CO2Australian Journal of Botany 40: 657-677.

Reddy, K.K., Davidonis, G.H., Johnson, A.S. and Vinyard, B.T.  1999.  Temperature regime and carbon dioxide enrichment alter cotton boll development and fiber properties.  Agronomy Journal 91: 851-858.

Reddy, K.R., Robana, R.R., Hodges, H.F., Liu, X.J. and McKinion, J.M.  1998.  Interactions of CO2 enrichment and temperature on cotton growth and leaf characteristics.  Environmental and Experimental Botany 39: 117-129.

Stuhlfauth, T. and Fock, H.P.  1990.  Effect of whole season CO2 enrichment on the cultivation of a medicinal plant, Digitalis lanataJournal of Agronomy and Crop Science 164: 168-173.

Taub, D.R., Seeman, J.R. and Coleman, J.S.  2000.  Growth in elevated CO2 protects photosynthesis against high-temperature damage.  Plant, Cell and Environment 23: 649-656.

Wurr, D.C.E., Edmondson, R.N. and Fellows, J.R.  2000.  Climate change: a response surface study of the effects of CO2 and temperature on the growth of French beans.  Journal of Agricultural Science 135: 379-387.

Zhu, J., Goldstein, G. and Bartholomew, D.P.  1999.  Gas exchange and carbon isotope composition of Ananas comosus in response to elevated CO2 and temperature.  Plant, Cell and Environment 22: 999-1007.

Ziska, L.H.  1998.  The influence of root zone temperature on photosynthetic acclimation to elevated carbon dioxide concentrations.  Annals of Botany 81: 717-721.