Plant and Soil 225: 213–225, 2000.

© 2000 Kluwer Academic Publishers. Printed in the Netherlands.

213

Changes in soil CO2 and O2 concentrations when radiata pine is grown in

competition with pasture or weeds and possible feedbacks with radiata
pine root growth and respiration

Roger Sands1,∗ , Putranto B. Nugroho1, David W.M. Leung2 , Osbert J. Sun3 and Peter W.
Clinton3
1 Schoolof Forestry, University of Canterbury, Private Bag 4800, Christchurch, New Zealand; 2 Department of
Plant and Microbial Sciences, University of Canterbury, Private Bag 4800, Christchurch, New Zealand and 3 New
Zealand Forest Research Institute, PO Box 29237, Christchurch, New Zealand

Received 27 January 2000. Accepted in revised form 28 June 2000

Key words: competition, radiata pine, root respiration, soil CO2 , soil O2

Abstract
This study examined the reciprocal effects of growing ryegrass, lotus and other weed species in competition with
radiata pine on soil CO2 and O2 concentrations and on the growth and root respiration of the radiata pine. Soil
O2 concentrations decreased and soil CO2 concentrations increased with increasing soil depth. Radiata pine plus
competing species slightly reduced soil O2 concentrations and markedly increased soil CO2 concentrations (up to
40 mmol mol−1 ) compared with radiata pine alone. The dry weights of shoots and roots, and the root respiration
rates of radiata pine grown with competing vegetation were much less than those for radiata pine alone. This
probably was not solely caused by competition for nutrients water or light since adequate water and nutrients
were supplied to all treatments and the radiata pine overtopped the competing vegetation. When radiata pine roots
were raised in NaHCO3 solutions equivalent to a range of CO2 concentrations, succinate dehydrogenase activity
(a metabolic indicator of mitochondrial respiration) and elongation rates of roots decreased as CO2 concentrations
increased from 0 to 40 mmol mol−1 . This suggests that the elevated CO2 concentrations found in the experiments
in soil was the cause, at least in part, of the reduced growth of radiata pine in competition with other species.

Introduction the balance between the rate of diffusion between soil

air and the air above the soil surface and the rates of
Radiata pine (Pinus radiata D. Don) is a promin- consumption of O2 or production of CO2 in respiration
ent plantation species in the Southern Hemisphere. by plant roots and microorganisms (Boot, 1993; Far-
It is also grown with pasture in agroforestry sys- rell et al., 1993; Russell, 1988). Consequently, the
tems. Radiata pine competes with weeds (plantations) concentration of O2 in the soil may be less, and the
and pastures (agroforestry systems) for water, light concentration of CO2 in the soil may be greater than
and nutrients. However, there is also the possibility in the atmosphere above the soil, and these differences
that the competitive relationships between radiata pine may become more pronounced with increasing soil
and competing vegetation could be influenced by the depth and with increasing biological activity in the soil
composition of the soil air. (Boot, 1993; Buyanovsky and Wagner, 1983; Castelle
The roots of competing species and associated soil and Galloway, 1990; Ivan and Kosian, 1987; John-
microorganisms are a sink for O2 and a source of CO2 . son et al., 1994; King et al., 1986; Magnusson, 1994;
The concentrations O2 and CO2 in the soil depend on Parada et al., 1983; Pyatt and Smith, 1983).
Low levels of soil O2 and/or high levels of soil
∗ Author for correspondence CO2 can inhibit tree growth and these effects may
214

be species-specific (Bouma et al., 1997). For ex- and each pot was placed in a tray containing 2.5 cm of
ample, a high CO2 concentration around the roots water to minimise gas leakage from the bottom of the
reduced growth of peas (Pisum sativum) and sun- pots.
flowers (Helianthus annus) (Stolwijk and Thimann, Radiata pine seeds were stratified at 4 ◦ C for two
1957), increased the growth of potatoes (Solanum weeks after which 3 seeds were directly sown into
tuberosum) (Arteca et al., 1979) but had no growth each pot. After 1 month, the seedlings were thinned
effect in beans (Phaseolus vulgaris) (Bouma et al., leaving only one healthy seedling per pot. After 6
1997). High CO2 concentrations around the roots months ryegrass was sown into the pots at a density of
also reduced the root respiration rate of Douglas-fir 700 seeds m−2 and lotus at a density of 500 seeds m−2 .
(Pseudotsuga menziesii) (Qi et al., 1994) and cactus Soil water was maintained at field capacity by an auto-
pear (Opuntia ficus-indica) (Nobel and Palta, 1989), mated drip irrigation system and 45 gm of the slow
but did not affect root respiration of either citrus (Cit- release fertiliser OsmocotePlus (nine months release)
rus volkameriana) or P. vulgaris (Bouma et al., 1997). was added to each pot.
It is possible, therefore, that plants in competition re- Gas sampling chambers were installed at 10 cm,
spond differently to O2 and/or CO2 in the soil air and 25 cm, and 40 cm from the surface of the pots. The
that this could result in the competitive advantage of chambers were made from 3 mm internal diameter alu-
one species over another. Even if there are no species minium tube. Each chamber was 17 cm in length, with
differences, a change in the composition of the soil two lines of 15 holes of 1 mm diameter with a spacing
air caused by competing species could have a uniform of 3 mm between lines and 8 mm within lines. The
negative impact. The possibility that the composition tubes were horizontally inserted into the soil through
of the soil air influences the growth of radiata pine the wall of the pot with holes facing downward, leav-
in competition with weeds or pastures was examined ing 2 cm outside the pot. The join between the tube
in this study. The objectives were to measure the dy- and the pot was sealed with epoxy resin and petro-
namics of O2 and CO2 in soil where radiata pine was leum jelly, and the outer tube openings were fitted
grown with or without competition and to assess the with rubber stoppers so that soil gas could later be
impact of this on the growth and physiology of the extracted. The gas samples were collected every two
radiata pine. weeks commencing from 4 weeks after the sowing
of the pasture seed. 10 mL of air from the sampling
chambers was extracted using a gas tight syringe and
Materials and methods stored in 2 × 5 mL vacutainer tubes. Concentrations
of O2 and CO2 in the gas samples were determined
Glass-house experiments using gas chromatography (HP 5890 Series II). The
gas chromatograph was equipped with a thermal con-
Two experiments were conducted in 1997 and 1998. ductivity detector and used 25-m × 0.53 mm Poraplot
The first experiment contained three treatments: bare Q and molecular sieve 5A columns. Nitrogen was used
soil, radiata pine grown alone, and radiata pine grown as the carrier gas. A Scott Specialty standard gas, with
with ryegrass (Lolium perenne L.). The second exper- CO2 , O2 and N2 concentrations of 2%, 24% and 74%
iment contained three treatments: radiata pine grown by volume respectively, was used for calibration.
alone, radiata pine grown with ryegrass and radiata The height and stem diameter (3 cm from soil sur-
pine grown with lotus (Lotus pedunculatus cav.). The face) of the radiata pine was measured each two weeks
experiments were arranged in a randomised complete commencing from the introduction of the ryegrass or
block design with a single tree plot and six blocks. lotus. The radiata pine was harvested 13 weeks after
They were conducted in a glass-house with day and the pasture seed was sown. The radiata pines were
night temperatures ranging from 10 ◦ C to 30 ◦ C and divided into needles, stem and roots. Roots for both
an average of 26/16 ◦ C (day/night) with an extended radiata pine and the pasture species were separated
photoperiod of 16 hours, where natural day length was into soil depths of 0–10 cm, 10–25 cm and 25–50 cm.
supplemented with 400W sodium lamps suspended All plant material was weighed after oven drying at
above the plants. Pots (PVC, 30 cm diameter, 60 cm 75 ◦ C for 62 h.
height) were filled with a sandy-loam soil (passed The excised root respiration rates of radiata pine
through 2 mm sieve) compacted to a bulk density of were measured at the time of harvesting using a Licor-
1.1 g cm−3 . The pots had 8 cm of gravel at the bottom 6200 Photosynthesis System with a modified sample
 215

chamber set in a closed configuration. Approximately collected. Thermocouple wires were inserted into the
1 gm fresh weight of fine roots (< 1 mm diameter soil using a thin metal rod to the same depths as the
non-woody secondary and tertiary laterals) from 0– gas sampling tubes. The temperatures were measured
10 cm and 10–25 cm depth, were collected, rinsed free at 100 cm distance from the radiata trees, located
from soil and organic matter, placed on tissue papers randomly. Soil samples were collected at 0–10 cm,
to absorb the water, and then placed in the chamber. 10–20 cm and 20–30 cm soil depths at a distance
Roots were allowed to stand for between 5 and 10 min of 100 cm from the trees using a 5 cm internal dia-
after which the changes of CO2 concentration were meter soil auger. Soil moisture content was measured
monitored every 2 min for 30 min. The roots did not gravimetrically by oven drying samples at 82o C for
dry out over this 40 minute period. The measurements 72 h.
were made when CO2 concentration was between 800
and 900 µmol mol−1 . A water bath maintained the Root elongation and succinate dehydrogenase activity
temperature of the chamber between 22 and 23 ◦ C.
Radiata pine seeds were germinated on wet filter pa-
Field experiment per and grown in distilled water until the radicles were
about 2 cm in length. The germinated seedlings were
This study was located at Dunsandel, a semi-arid site grown in a controlled environment (80% relative hu-
on the Canterbury Plains of New Zealand. The experi- midity, 700 µmol m−2 s−1 PPFD for 18 h a day) at
ment was conducted in part of a radiata pine and weed temperatures of 15, 20 and 25 ◦ C in 20 litre plastic
competition study established in 1996 (see Mason and containers containing NaHCO3 solutions equivalent
Kirongo, 1999). The soil type was a Lismore stony to 0, 10, 20 and 40 mmol mol−1 CO2 concentra-
silt loam; the average annual rainfall was 600 mm, tions. The roots of intact seedlings (60 per container)
occurring predominantly in winter. The understorey were inserted through holes in the lid so that the cole-
species were Italian ryegrass (Lolium multiflorum) and optiles were outside the solution while the roots were
clover (Trifolium repens), with some sorrel (Rumex submerged in the solutions. A reusable putty (Blue-
acetosella). Sampling sites were selected from Block tak, Bostik New Zealand) was used to seal the gap
3 (Mason and Kirongo, 1999) in which weeds present between the holes and the roots. The solutions were
(with-weeds) and weeds absent (without-weeds) treat- prepared by diluting NaHCO3 in 0.001 M sodium
ments were adjacent. Within each treatment area (ap- phosphate buffer at pH 7. Using Henry’s Law and
proximately 36 × 27 m2 ), there were 70 radiata pine the Henderson-Hasselbach equation (Umbreit et al.,
trees consisting of 7 clones, arranged in one line per 1972), the concentrations of NaHCO3 equivalent to
clone. The planting distance was 3 × 3 m. Five sample 1%, 2% and 4% CO2 concentrations at 15 ◦ C, 20 ◦ C
trees from clone 4 (located on the same planting line) and 25 ◦ C are (1.778 mM, 3.556 mM, 7.111 mM),
within each treatment area were selected randomly. (1.589 mM, 3.178 mM, 6.357 mM) and (1.462 mM,
The radiata pines and the weeding treatments had been 2.924 mM, 5.848 mM), respectively. For each exper-
established for 3 years. iment, the sodium phosphate buffer without NaHCO3
A tube (3 mm internal diameter aluminium) was was used as a 0% of CO2 concentration control. The
inserted vertically into the soil at each of 10 cm, 20 cm solutions were renewed every 24 h. Seedlings were
and 30 cm soil depths at a distance of 25 cm from also grown at 20 ◦ C in 7.16 mM NaCl under the same
each tree. The tubes were open at both ends, with the growing environment to determine whether there was
lower 2 cm containing 16 equally spaced holes drilled a salinity effect at concentrations of NaCl equivalent
through the tubing. The tops of the access tubes exten- to the highest concentration of NaHCO3 .
ded about 5 cm above the soil surface and were capped Root elongation was measured on 20 randomly
with a rubber stopper. Gas samples were collected by chosen seedlings per treatment at 1, 2, 3, 4, 7 and
extracting 10 mL gas samples using a gas tight syr- 14 days. After 14 days, the seedlings were harves-
inge through the rubber stopper and stored in 2 × 5 ted. Root segments (10 mm length from the root tip)
mL vacutainer tubes. Concentrations of O2 and CO2 from each container were bulked and stored in liquid
were determined by gas chromatography in a manner N2 prior to measurement of succinate dehydrogenase
similar to the glass-house experiment. activity. Succinate dehydrogenase activity was only
Soil temperature was measured with a hand-held measured on seedlings grown at 25 ◦ C. Frozen root
thermocouple thermometer when the soil gases were material was ground in a mortar and pestle with liquid
 216

Figure 1. Soil O2 concentrations at 10, 25 and 40 cm soil depth from glasshouse experiments 1 and 2 (± S.E., n=6).
Figure 2. Soil CO2 concentrations at 10, 25 and 40 cm soil depth from glasshouse experiments 1 and 2 (±S.E., n=6).
217
218

Figure 3. Biomass distribution of radiata pine from glasshouse experiments 1 and 2. Means between treatments within a biomass component
with different letters were significantly different at p<0.05 (± S.E., n=6).

nitrogen. Buffer (50 mM sodium phosphate buffer at grown with lotus were similar or slightly higher than
pH7) was added to the root tissue substrate (1 ml of concentrations under radiata pine without competing
buffer to 50 mg fresh weight of root tissue) and the vegetation. However after the third sampling, soil O2
homogenate was centrifuged at 4 ◦ C for 20 min at concentrations decreased (Figure 1). The concentra-
13,500 rpm. The supernatant was added to 50 mM tion of soil O2 ranged from 175 to 209 mmol mol−1
sodium phosphate buffer (pH 7) supplemented with in Experiment 1 and from 184 to 209 mmol mol−1
0.1% tetrazolium chloride and 0.01 M sodium succin- in Experiment 2. These concentrations were slightly
ate. After 60 minutes incubation at 37 ◦ C the reaction lower than those found in the atmosphere (about 210
was terminated by boiling for 5 minutes then adding mmol mol−1 ).
1 N NaOH. The precipitate was collected by cent- In both experiments, soil CO2 concentrations in-
rifuging for 15 min and it was then dissolved with creased with increasing soil depth and time (Figure 2).
C2 H5 OH. Absorbance was measured at 494 nm. Suc- Soil CO2 concentrations under radiata pine grown
cinate dehydrogenase activity was measured as the without competing vegetation, were lower than where
rate of reduction of tetrazolium chloride calculated radiata pine was grown with either ryegrass (Figure 2,
using an equation generated from the standard curve. Experiments 1 and 2) or lotus (Figure 2, Experiment
2). The effect of lotus changed with time. For the
first three sampling occasions, soil CO2 concentra-
Results tions under radiata pine grown with lotus were lower
than radiata pine grown with ryegrass. Later, soil
Glass-house experiment CO2 concentrations under radiata pine grown with
lotus steadily increased and exceeded concentrations
In both experiments, soil O2 concentration decreased under radiata pine grown with ryegrass (Figure 2, Ex-
with increasing soil depth and with increasing time periment 2). Soil CO2 concentrations ranged from
(Figure 1). The average soil O2 concentration under 2 to 45 mmol mol−1 in Experiment 1 and from 2
radiata pine grown with ryegrass was lower than that to 25 mmol mol−1 in Experiment 2. These concen-
under radiata pine grown with lotus (Figure 1, Ex- trations were more than 100 times higher than the
periment 2) and lower than under radiata pine grown CO2 concentration in the atmosphere (about 350 µmol
without competing vegetation (Figure 1, Experiments mol−1 ).
1 and 2). The effect of lotus changed with time. The biomass of radiata pine grown without com-
Initially, soil O2 concentrations under radiata pine peting vegetation was larger than either pine grown
 219
Table 1. Dry weight (g) of roots of radiata pine, ryegrass and lotus
with lotus (Figure 3, Experiment 2) or pine grown when radiata pine was growing alone or with ryegrass in Experiment
with ryegrass (Figure 3, Experiments 1 and 2). The 1, and when radiata pine was growing alone, with ryegrass or with
total biomass of radiata pine without competing ve- lotus in Experiment 2. Means between treatments within an exper-
getation was nearly twice that of pine grown with iment with different letters are significantly different at p<0.05 (±
S.E., n=6)
ryegrass. This pattern was similar for each compon-
ent of the radiata pine although the difference between Radiata pine alone Radiata pine + Radiata pine +
the root dry weight of radiata pine grown with lotus ryegrass lotus
and radiata pine grown with ryegrass was not signi-
ficant. Even though analysis of variance showed that Experiment 1
radiata pine 44.6a ± 2.3 26.2b ± 3.2 –
there were no significant differences (p>0.05) in the
ryegrass – 127.9c ± 25.3 –
root/shoot ratio in Experiment 1 (mean 0.45) and Ex-
periment 2 (mean 0.40), the mean value for radiata
Experiment 2
pine with ryegrass (0.52) was noticeably higher than
radiata pine 30.0a ± 3.4 22.5b ± 3.5 23.3b ± 4.0
that of radiata pine with lotus (0.37) and radiata pine ryegrass – 63.5c ± 14.0
alone (0.32). The root biomass of radiata pine grown lotus – – 28.1ab ± 3.9
with competing vegetation was lower than when radi-
ata pine was grown without competing vegetation in
both experiments (Figure 3 and Table 1).
The total root biomass (radiata pine roots plus roots Table 2. Root respiration rates of radiata pine growing alone,
of competing vegetation) in the soil under radiata pine with ryegrass and with lotus in Experiments 1 and 2. Values
grown with competing vegetation was far greater than with different letters are significantly different at p<0.05 (±S.E.,
n=6)
where radiata pine was grown without competing ve-
getation (Table 1). The effect of soil depth on root Radiata pine root respiration rates
biomass was significant (p<0.05). In both experiments (nmol g−1 s−1 )
42–46% of the radiata pine root mass was located at Experiment 1 Experiment 2
the 0–10 cm depth, 28–31% at 10–25 cm depth and
Radiata pine alone 12.5a ± 1.25 12.6a ± 2.8
24–28% in the lowest part of the pot. The distribu-
radiata pine + ryegrass 5.6b ± 0.47 6.0b ± 1.3
tion of the ryegrass roots in the radiata pine-ryegrass
radiata pine + lotus 7.5b ± 1.7
treatment was similar to that of the radiata pine; root
mass was 41%, at 0–10 cm, 27% at 10–25cm and 32%
at 25–50 cm. The lotus root biomass was distributed
more often in the top part of the soil, where there was
approximately 60% at 0–10 cm, 29% at 10–25 cm and consistently lower than where radiata pine was grown
31% at 25-50 cm depth. without weeds. At the same depth, soil CO2 concen-
There were no significant differences (p<0.05) trations under the radiata pine with weeds were nearly
in root respiration rates between 10 cm and 25 cm twice those under radiata pine without weeds (Fig-
soil depths. The root respiration rate of radiata pine ure 4). The highest soil CO2 concentration found in
grown without competing vegetation was about twice the with-weeds treatment was 6.1 mmol mol−1 and
that of radiata pine grown with competing vegetation in the without-weeds treatment was 4.2 mmol mol−1 .
(Table 2). The lowest soil O2 concentrations in the with-weeds
treatment was 182 mmol mol−1 and in the without-
Field experiment weeds treatment, 195 mmol mol−1 (Figure 4). In both
treatments, the maximum soil CO2 and minimum soil
Figure 4 shows soil CO2 concentration (A), O2 con- O2 concentrations were found in February, which co-
centration (B), temperature (C) and moisture content incided with peak soil temperature and the lowest soil
(D) at various times in the field experiment. There moisture content (Figure 4). The elevation of soil CO2
was an increase in CO2 concentrations and a decrease concentrations and depression of soil O2 concentra-
in O2 concentrations with increasing soil depth (Fig- tions were not as great in the field trial (Figure 4) as in
ure 4). At each sampling time and depth, soil CO2 the glasshouse experiments (Figures 1 and 2).
concentrations under radiata pine with weeds were Soil temperatures under the without-weeds treat-
consistently higher and soil O2 concentrations were ment were consistently higher than under the with-
220

Figure 4. Soil CO2 concentrations (A), O2 concentrations (B), temperatures (C) and moisture contents (D) at 10, 20 and 30 cm soil depth in
the radiata pine field experiment with and without weeds (± S.E., n=5).
 221

Figure 5. Relationship between soil temperature and soil CO2 concentration at 10, 20 and 30 cm soil depth under radiata pine with weeds (2)
and without weeds (N). Slopes between treatments were significantly different at p<0.05.
222

Figure 6. Root elongation of radiata pine during 14 days in NaHCO3 rooting medium after treatment with different carbon dioxide
concentrations (± S.E., each point is the mean of 3 temperatures × 20 replicates).

weeds treatment (Figure 4). The differences were

greater nearer the soil surface. The greatest difference
was approximately 4 ◦ C during February 1999, when
the highest soil temperature was recorded. On each
sampling occasion, soil moisture content at both 20
and 30 cm depths under the with-weeds treatment was
consistently lower than in the without-weeds treat-
ment. This consistency was not found at the 10 cm
depth (Figure 4).
No significant correlation was found between
either soil O2 concentration and soil temperature, or
soil O2 and soil moisture content (p>0.05). There was
a significant correlation (p<0.05) between soil CO2
concentration and temperature in both treatments at
each depth (r values from 0.62 to 0.84). Soil CO2
concentrations increased with increasing soil temper-
ature and the rate of increase was greater when the
trees were in competition with weeds (Figure 5). How-
ever, soil moisture content had a weaker relationship
with soil CO2 concentration. A significant correlation
(p<0.05) was found in the with-weeds treatment, at
both 20 cm and 30 cm depths (r = -0.59). Figure 7. Reduced tetrazolium chloride in radiata pine root tissue
as a function of CO2 concentration in the rooting medium (±S.E.,
n=4).
 223

Root elongation rate and succinate dehydrogenase at concentrations of oxygen in soil air of <100 mmol
activity mol−1 . However, even in the well-aerated soil used in
this study (air-filled porosity >20% at –0.01 MPa soil
Root elongation of radiata pine in the experiments in matric potential) the increase in the concentration of
which radiata pine seedlings were grown in NaHCO3 CO2 was substantial, up to 40 mmol mol−1 which is
solutions was reduced as CO2 concentration increased >100 × that in the atmosphere and 20 × that in the
(Figure 6). Compared with the control treatment, the bare soil at 15 cm soil depth (Figure 2). Concentra-
average inhibition of root elongation at 14 days was tions of CO2 in soil as low as 1.5 mmol mol−1 have
14.4, 20.3 and 38.1%, for 10, 20 and 40 mmol mol−1 been shown to reduce the respiration rate of roots of
CO2 concentration respectively. There was no ef- Pinus radiata (Ryan et al., 1996) and Acer saccharum
fect of 7.16 mM NaCl on root elongation rate (data (Burton et al., 1997). Therefore the emphasis has been
not shown) and there was no significant effect of on CO2 rather than O2 in this study.
temperature. It is well known that an increase in vegetation
The succinate dehydrogenase enzyme activ- density can be associated with an increase in soil
ity of the root tissue decreased with increasing CO2 (Boot, 1993; Magnusson, 1992; Parada et al.,
CO2 concentration (Figure 7). At 10 mmol mol−1 1983). For example, in the Kielder forest, King et
CO2 , the activity of succinate dehydrogenase was 72% al., (1986) found higher soil CO2 concentrations under
that of the control (0 mmol mol−1 CO2 ). At 20 mmol lodgepole pine with a grass understorey than where
mol−1 CO2 the activity was 56% of the control, and at lodgepole pines were grown alone. Clear-felled forests
40 mmol mol−1 CO2 it was 59% of the control. have lower soil CO2 concentrations than forested areas
(Boot 1993) and the highest soil CO2 concentration in
Arizona soils occurs in the most densely vegetated
Discussion soils (Magnusson, 1992). In the glasshouse experi-
ment in this study, the higher soil CO2 concentrations
The concentration of O2 and CO2 at any point in the for radiata pine grown with either ryegrass or lotus
soil is a result of a combination of the physical prop- (Figure 2) were probably due to the greater biomass
erties of the soil that determine the rate of gaseous of respiring root (Table 1). This is probably also true
exchange with the atmosphere, as well as live roots of the field experiment (Figure 4), although there were
and microorganisms which use O2 and discharge CO2 no supporting measurements of total root biomass in
in respiration. In this study the presence of living roots this trial.
with associated microbial activity decreased O2 con- Soil CO2 concentrations in the field experiment
centrations and increased CO2 concentrations in the were strongly correlated with soil temperature and
soil (Figures 1, 2 and 4). Also, O2 concentrations de- poorly correlated with moisture content. This is con-
creased and CO2 concentrations increased with soil sistent with other studies of soil CO2 (Garret and
depth (Figures 1, 2 and 4) because atmospheric ex- Cox, 1973; Gunn and Trudgill, 1982; Kowalenko et
change of these gases becomes more difficult with al., 1978). Soil CO2 concentrations increased with
increasing soil depth. increasing soil temperature and this rate of increase
Radiata pine growing with competing species was was greater when radiata pine was in competition with
associated with a decrease in soil O2 concentrations weeds (Figure 5). However, the soils in the with-
and an increase in CO2 concentrations, when com- weeds treatment had lower temperatures than in the
pared with radiata pine grown alone (Figures 1, 2 and without-weeds treatment (Figure 4).
4). The decrease in O2 concentrations, though signi- Radiata pine competing with ryegrass or lotus in
ficant, was small and unlikely to be associated with the glasshouse experiments were slower growing than
growth inhibition. Oxygen concentrations were never radiata pine grown alone (Figure 3 and Table 1).
below 175 mmol mol−1 . The literature suggests that Competition for water and nutrients by competing
growth inhibition occurs at much lower O2 concen- vegetation is known to reduce the growth of radi-
trations (Crawford and Braendle, 1996; Drew, 1992; ata pine (Clinton, 1990; Mead and Mansur, 1993).
Kriedemann and Sands, 1984) characteristic of hy- However, the glasshouse experiment was kept well
dric or near hydric soils (Faulkner and Patrick, 1992). watered and well supplied with nutrients. It is unlikely
Kramer and Boyer (1995) considered significant ef- that competition for water and/or nutrients was the
fects on growth of oxygen deprivation occured only sole factor in this instance. Usually, if water and/or
224

nutrients are limiting, a plant will preferentially alloc- slowly from another cause which is independent of
ate assimilate to roots (Sands and Mulligan, 1989). soil CO2 concentrations. However, in the experi-
There were no significant treatment effects on the ments with radiata pine roots growing in solutions of
root/shoot ratio in this study which suggests that wa- NaHCO3 , it was shown that increasing concentrations
ter and/or nutrients were not dominating factors and of CO2 in the rooting medium both reduced the rate
that other factor(s) might be contributing to the the of root elongation (Figure 6) and succinate dehyd-
growth inhibition of radiata pine growing with com- rogenase activity (Figure 7) over the same range of
peting species. The root/shoot ratio of the pine plus CO2 concentrations found in the glasshouse and field
ryegrass was greater (though not significantly) in Ex- trials (Figures 2 and 4). (The fact that NaCl at the
periment 2 which suggests that water and/or nutrition same concentration as the highest NaHCO3 concen-
might have been more important in this treatment. tration did not affect root elongation rate suggests that
The radiata pine overtopped the ryegrass and lotus there was no complicating salinity effect in the use of
and it is therefore unlikely that competition for light NaHCO3 solutions in this study). This is in accord
was a factor. In the field experiment, soil temperat- with Gonzàles-Meler et al. (1996) who showed that
ure in the without-weeds treatment was higher than an increase of 365 µmol mol−1 CO2 concentration
the with-weeds treatment (Figure 4). These higher inhibited the activity of succinate dehydrogenase in
soil temperatures could favour the growth of the trees soybean mitochondria. Succinate dehydrogenase is a
without weeds. However, no significant differences key enzyme in respiration. In this study the succinate
in soil temperatures were found in the glasshouse dehydrogenase activity of radiata pine roots was re-
experiment (data not shown); therefore, soil temperat- duced to about 60% of that in the no CO2 treatment
ure differences cannot be contributing to competitive in 20–40 mmol mol−1 concentrations of CO2 in the
effects in this instance. rooting medium. Ryan et al. (1996) estimated that ra-
Perhaps the concentration of CO2 in the soil air had diata pine root respiration at approximately 1.5 mmol
an effect. The glasshouse experiment clearly showed mol−1 soil CO2 concentration was only 23% of that
that the presence of ryegrass and lotus under radiata at 0.4 mmol mol−1 soil CO2 concentration. The dif-
pine increased soil CO2 concentrations and inhibited ference in the level of inhibition between the studies
the growth and root respiration rate of the radiata may be due to factors other than succinate dehyd-
pine. A negative effect of high CO2 concentration on rogenase activity contributing to the inhibitory effect
growth has been reported for a range of plant species of CO2 concentration on the respiration rate of roots
(Stolwijk and Thimann, 1957) including tree species (Gonzàles-Meler and Siedow, 1999).
(Leyton and Rousseau, 1957; McDowell et al., 1999; It is, therefore, likely that the increased CO2 con-
Nobel and Palta, 1989; Palta and Nobel, 1989; Qi centrations in the glasshouse experiment contributed
et al., 1994). The root respiration rate of Pinus ra- to the inhibition of the growth and root respiration rate
diata D. Don at 1.5 mmol mol−1 CO2 was found to of radiata pine in competition with ryegrass or lotus. It
be only 23% of that measured at 0. 4 mmol mol−1 is known that soil CO2 concentrations can have differ-
CO2 concentration (Ryan et al., 1996), and that of ent effects on different species (see Introduction). This
Douglas fir decreased 4–5 times when CO2 concen- raises the intriguing and as yet untested possibility
tration doubled (Qi et al. 1994). Root respiration of that the CO2 concentration of the soil atmosphere may
sugar maple (Acer saccharum) was sensitive below 1.5 play an important role in the competitive relationships
mmol mol−1 CO2 in the rooting medium but stable between plants.
at above 6 mmol mol−1 CO2 (Burton et al., 1997).
However, Bouma et al. (1997) observed citrus (Citrus
volcamerica) rootrespiration was independent of CO2
concentration over the range 0 to 25 mmol mol−1 . Acknowledgements
It has not though been proven in the glasshouse
experiment that the increased soil CO2 concentrations We thank Irene Hudson for advice on statistics and
were directly responsible for the inhibited growth and David Macpherson, Dave Condor, Vicky Wilton and
root respiration rate of the radiata pine in compet- Bob Bullsmith for technical assistance.
ition with ryegrass or lotus. It is possible that the
reduced respiration rate of the radiata pine roots was
the consequence of the radiata pine growing more
 225

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