Global Change Biology (1999) 5, 255–267
Monoterpene emission from coniferous trees in response
to elevated CO
2
concentration and climate warming
JOHN V. H. CONSTABLE*, MARCY E. LITVAK*, JAMES P. GREENBERG† and
RUSSELL K. MONSON*
*Department of EPO Biology, University of Colorado, Boulder, CO 80309–0334, USA, †Atmospheric Chemistry Division,
National Center for Atmospheric Research, Boulder, CO 80307, USA
Abstract
It was hypothesized that high CO
2
availability would increase monoterpene emission
to the atmosphere. This hypothesis was based on resource allocation theory which
predicts increased production of plant secondary compounds when carbon is in excess
of that required for growth. Monoterpene emission rates were measured from needles
of (a) Ponderosa pine grown at different CO
2
concentrations and soil nitrogen levels,
and (b) Douglas fir grown at different CO
2
concentrations. Ponderosa pine grown at
700 μmol mol
–1
CO
2
exhibited increased photosynthetic rates and needle starch to
nitrogen (N) ratios when compared to trees grown at 350 μmol mol
–1
CO
2
. Nitrogen
availability had no consistent effect on photosynthesis. Douglas fir grown at
550 μmol mol
–1
CO
2
exhibited increased photosynthetic rates as compared to growth at
350 μmol mol
–1
CO
2
in old, but not young needles, and there was no influence on the
starch/N ratio. In neither species was there a significant effect of elevated growth CO
2
on needle monoterpene concentration or emission rate. The influence of climate warming
and leaf area index (LAI) on monoterpene emission were also investigated. Douglas fir
grown at elevated CO
2
plus a 4 °C increase in growth temperature exhibited no change
in needle monoterpene concentration, despite a predicted 50% increase in emission rate.
At elevated CO
2
concentration the LAI increased in Ponderosa pine, but not Douglas
fir. The combination of increased LAI and climate warming are predicted to cause an
80% increase in monoterpene emissions from Ponderosa pine forests and a 50% increase
in emissions from Douglas fir forests. This study demonstrates that although growth at
elevated CO
2
may not affect the rate of monoterpene emission per unit biomass, the
effect of elevated CO
2
on LAI, and the effect of climate warming on monoterpene
biosynthesis and volatilization, could increase canopy monoterpene emission rate.
Keywords: atmospheric chemistry, carbon dioxide, climate change, nitrogen, Pinus ponderosa,
Pseudotsuga menziesii
Received 27 October 1997; revised version received 27 January and accepted 7 February 1998
Introduction
The emission of volatile organic compounds (VOCs) from
vegetation exerts the dominant control over regional
oxidative chemistry of the atmosphere in many terrestrial
ecosystems (Monson et al. 1995; Lerdau et al. 1997). In
the case of coniferous forests, monoterpene compounds
represent the dominant biogenic VOC (Lerdau 1991).
Monoterpenes are 10-carbon compounds used primarily
for herbivore defence, and volatilization to the atmo-
Correspondence: John V.H. Constable, Department of Biological
Sciences, University of Illinois at Chicago, 845 West Taylor
St., Chicago, IL 60607, USA, fax +1/312-996-2805, e-mail
constabj@tigger.cc.ulc.edu
© 1999 Blackwell Science Ltd. 255
sphere occurs from storage reservoirs in the needles and
bark of conifers. Once in the atmosphere, monoterpenes
react readily with oxidative species (e.g. hydroxyl radical,
nitrate radical, and ozone) to form a variety of secondary
products that influence atmospheric chemistry and cli-
mate change (Wuebbles et al. 1989). Among the products
of monoterpene oxidation are CO, organic acids, organic
nitrates, and reactive organic peroxy compounds
(Fehsenfeld et al. 1992). Thus, monoterpene emissions
from coniferous forests represent a primary contribution
to the broader discipline of biosphere–atmosphere inter-
actions.
Theoretical models of resource allocation suggest that