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Trees (Types - Pine) - Summary
Nearly all woody species respond to increases in the air's CO2 content by displaying enhanced rates of photosynthesis and biomass production.  In this summary, we review several recently published responses of pine (genus Pinus) trees to atmospheric CO2 enrichment.

In the relatively short-term study of Tjoelker et al. (1998a), Jack pine seedlings grown for three months at atmospheric CO2 concentrations of 580 ppm exhibited photosynthetic rates that were about 28% greater than those displayed by control seedlings fumigated with air containing 370 ppm CO2.  Similarly, Hymus et al. (1999) reported that a 210-ppm increase in the air's CO2 content boosted rates of net photosynthesis in 15-year-old loblolly pine trees by 65% in the warmer months of an annual photosynthetic study.  In a pair of two-year studies, twice-ambient concentrations of atmospheric CO2 boosted rates of photosynthesis in seedlings of ponderosa and longleaf pine by 49% (Houpis et al., 1999) and 50% (Runion et al., 1999), respectively.  In the three-year study of Kainulainen et al. (1998), Scots pine trees simultaneously exposed to elevated CO2 and elevated O3 concentrations exhibited no photosynthetically-derived increases in starch production until the final year of the study.  Last of all, at the conclusion of four years of treatment exposure, Turnbull et al. (1998) reported that Pinus radiata seedlings grown at 650 ppm CO2 displayed photosynthetic rates that were 63% greater than rates displayed by control seedlings growing at 360 ppm CO2.

Because elevated CO2 enhances photosynthetic rates in pine trees, it likely also leads to increased biomass production in these important coniferous trees.  And so it does.  In the short-term three-month study of Tjoelker et al. (1998b), for example, Jack pine seedlings receiving an extra 210 ppm CO2 displayed final dry weights that were about 20% greater than those displayed by seedlings growing at ambient CO2 concentrations.  Similarly, after growing loblolly pine seedlings for four months with an extra 300 ppm of CO2, Gavazzi et al. (2000) noted that elevated CO2 enhanced seedling biomass by 22%.  In the six-month study of Janssens et al. (1998), twice-ambient CO2 concentrations increased root dry mass in Scots pine seedlings by 135%.  And in another six-month study, Maherali and DeLucia (2000) reported that a 750-ppm increase in the CO2 content of the air increased the biomass of ponderosa pines by 42 and 62% at low and high air temperatures, respectively.

Similar increases in pine seedling biomass have been reported for CO2 enrichment studies of longer duration.  After the first year of treatment exposure in a FACE experiment, for example, loblolly pine trees exposed to an additional 210 ppm of CO2 displayed initial biomass values that were 14% greater than those exhibited by their ambiently-grown counterparts (Naidu and DeLucia, 1999).  Likewise, in the 1.5-year study of Entry et al. (1998), longleaf pine grown at 720 ppm CO2 attained whole-plant biomass values that were 42% greater than those attained by seedlings grown at 365 ppm CO2.  Moving on to the two-year study of Walker et al. (1998a), it was reported that a doubling of the atmospheric CO2 concentration increased root and shoot dry weights by about 84%.  Finally, in the four-year study of Tissue et al. (1997), a 300-ppm increase in the atmospheric CO2 concentration increased seedling biomass by 90%.  In addition, in a literature review of their own previously published research, Johnson et al. (1998) note that twice-ambient levels of atmospheric CO2 can enhance ponderosa and loblolly pine seedling biomass by as much as 1000% under conditions of moderate soil nitrogen deficiency.

Elevated CO2 has also been documented to improve root colonization in pine trees by various species of ectomycorrhizal fungi, which enhance nutrient and water acquisition for their symbiotic hosts.  After just four months exposure to twice-ambient concentrations of atmospheric CO2, for example, Rouhier and Reed (1998) reported that the number of fungal hyphal tips associated with roots of Scots pine had increased by 62%.  Similarly, a doubling of the atmospheric CO2 concentration increased the number of root tips in eastern white pine seedlings by 49%, which allowed 38% more root area to become colonized by ectomycorrhizal fungi (Godbold et al., 1997).  Likewise, after one year of differential treatment exposure, Walker et al. (1998b) reported that the total number of ectomycorrhizal fungi associated with ponderosa pine seedling roots grown at 700 ppm CO2 was 85% greater than that associated with roots of seedlings grown at 350 ppm CO2.

Finally, in a somewhat different type of study, Scherzel et al. (1998) reported that twice-ambient atmospheric CO2 concentrations had no effect on the decomposition of eastern white pine seedling litter.

In summary, it is clear that as the CO2 content of the air increases, pine trees will likely display enhanced rates of photosynthesis and biomass production, resulting in greater carbon sequestration by this coniferous genus.  In addition, as the air's CO2 content increases, it is likely that greater root colonization by ectomycorrhizal fungi will occur, which may further increase biomass production in these woody species.

For more information on pine growth responses to atmospheric CO2 enrichment see Plant Growth Data: Eastern White Pine (dry weight), Loblolly Pine (dry weight, photosynthesis), Longleaf Pine (dry weight, photosynthesis), Monterey Pine (photosynthesis), Ponderosa Pine (dry weight, photosynthesis), and Scots Pine (dry weight, photosynthesis).

References
Entry, J.A., Runion, G.B., Prior, S.A., Mitchell, R.J. and Rogers, H.H.  1998.  Influence of CO2 enrichment and nitrogen fertilization on tissue chemistry and carbon allocation in longleaf pine seedlings.  Plant and Soil 200: 3-11.

Gavazzi, M., Seiler, J., Aust, W. and Zedaker, S.  2000.  The influence of elevated carbon dioxide and water availability on herbaceous weed development and growth of transplanted loblolly pine (Pinus taeda).  Environmental and Experimental Botany 44: 185-194.

Godbold, D.L., Berntson, G.M. and Bazzaz, F.A.  1997.  Growth and mycorrhizal colonization of three North American tress species under elevated atmospheric CO2New Phytologist 137: 433-440.

Houpis, J.L.J., Anderson, P.D., Pushnik, J.C. and Anschel, D.J.  1999.  Among-provenance variability of gas exchange and growth in response to long-term elevated CO2 exposure.  Water, Air, and Soil Pollution 116: 403-412.

Hymus, G.J., Ellsworth, D.S., Baker, N.R. and Long, S.P.  1999.  Does free-air carbon dioxide enrichment affect photochemical energy use by evergreen trees in different seasons?  A chlorophyll fluorescence study of mature loblolly pine.  Plant Physiology 120: 1183-1191.

Janssens, I.A., Crookshanks, M., Taylor, G. and Ceulemans, R.  1998.  Elevated atmospheric CO2 increases fine root production, respiration, rhizosphere respiration and soil CO2 efflux in Scots pine seedlings.  Global Change Biology 4: 871-878.

Johnson, D.W., Thomas, R.B., Griffin, K.L., Tissue, D.T., Ball, J.T., Strain, B.R. and Walker, R.F.  1998.  Effects of carbon dioxide and nitrogen on growth and nitrogen uptake in ponderosa and loblolly pine.  Journal of Environmental Quality 27: 414-425.

Kainulainen, P., Holopainen, J.K. and Holopainen, T.  1998.  The influence of elevated CO2 and O3 concentrations on Scots pine needles: Changes in starch and secondary metabolites over three exposure years.  Oecologia 114: 455-460.

Maherali, H. and DeLucia, E.H.  2000.  Interactive effects of elevated CO2 and temperature on water transport in ponderosa pine.  American Journal of Botany 87: 243-249.

Naidu, S.L. and Delucia, E.H.  1999.  First-year growth response of trees in an intact forest exposed to elevated CO2Global Change Biology 5: 609-613.

Rouhier, H. and Read, D.J.  1998.  Plant and fungal responses to elevated atmospheric carbon dioxide in mycorrhizal seedlings of Pinus sylvestrisEnvironmental and Experimental Botany 40: 237-246.

Runion, G.B., Mitchell, R.J., Green, T.H., Prior, S.A., Rogers, H.H. and Gjerstad, D.H.  1999.  Longleaf pine photosynthetic response to soil resource availability and elevated atmospheric carbon dioxide.  Journal of Environmental Quality 28: 880-887.

Scherzel, A.J., Rebbeck, J. and Boerner, R.E.J.  1998.  Foliar nitrogen dynamics and decomposition of yellow-poplar and eastern white pine during four seasons of exposure to elevated ozone and carbon dioxide.  Forest Ecology and Management 109: 355-366.

Tissue, D.T., Thomas, R.B. and Strain, B.R.  1997.  Atmospheric CO2 enrichment increases growth and photosynthesis of Pinus taeda: a 4 year experiment in the field.  Plant, Cell and Environment 20: 1123-1134.

Tjoelker, M.G., Oleksyn, J. and Reich, P.B.  1998a.  Seedlings of five boreal tree species differ in acclimation of net photosynthesis to elevated CO2 and temperature.  Tree Physiology 18: 715-726.

Tjoelker, M.G., Oleksyn, J. and Reich, P.B.  1998b.  Temperature and ontogeny mediate growth response to elevated CO2 in seedlings of five boreal tree species.  New Phytologist 140: 197-210.

Turnbull, M.H., Tissue, D.T., Griffin, K.L., Rogers, G.N.D. and Whitehead, D.  1998.  Photosynthetic acclimation to long-term exposure to elevated CO2 concentration in Pinus radiata D. Don. is related to age of needles.  Plant, Cell and Environment 21: 1019-1028.

Walker, R.F., Geisinger, D.R., Johnson, D.W. and Ball, J.T.  1998a.  Atmospheric CO2 enrichment and soil N fertility effects on juvenile ponderosa pine: Growth, ectomycorrhizal development, and xylem water potential.  Forest Ecology and Management 102: 33-44.

Walker, R.F., Johnson, D.W., Geisinger, D.R. and Ball, J.T.  1998b.  Growth and ectomycorrhizal colonization of ponderosa pine seedlings supplied different levels of atmospheric CO2 and soil N and P.  Forest Ecology and Management 109: 9-20.