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Abstract
The response of potato (Solanum tuberosum L., cv. Shepody) to elevated CO2 was studied in 1994 at Fairbanks, Alaska using open-top
chambers (OTC). Three CO2 exposure levels were used in OTC: ambient (A); A + 175 mmol mol1 CO2 (A + 175); A + 350 mmol mol1
CO2 (A + 350). Plots were also established outside chambers (ANC) to test chamber affects. Potato leaf appearance rate and canopy
development did not differ between CO2 treatments. Six plants in each plot were harvested in late July and August. At the first harvest, the
number of stems, total stem weight, number of flowers, and total flower dry weight decreased with increasing CO2. Biomass allocated to stems
and leaves declined while allocation to tubers increased with elevated CO2. Root:shoot ratio was larger and percent leaf nitrogen declined
12.6% with increased CO2. At the final harvest, total tuber dry weight in A + 350 chambers was 36% higher than in the A treatment, but yields
in open-top chamber plots were lower than the no-chamber plots possibly due to 714% less photosynthetic active radiation in chambers. Net
photosynthesis measurements made on July 13 showed no differences due to CO2 treatment; however, on August 5, net photosynthesis of
leaves grown at A + 350 was 53% greater than at ambient CO2. Tuber initiation and growth began between the two photosynthesis
measurements. Before tuber initiation, photosynthetic rates may have been down-regulated by insufficient carbon sink capacity.
# 2005 Elsevier B.V. All rights reserved.
Keywords: Carbon dioxide; Atmospheric CO2; Potato; Yield; Photosynthesis; Open-top chambers; Subarctic
1. Introduction
Kimball (1983) compiled data from 430 studies with
various crop species grown under ambient and 2X ambient
CO2 levels and found that yields at 2X CO2 were increased
on average by 32% over ambient. However, lack of response
to elevated atmospheric CO2 or apparent acclimation has
been noted in a number of studies. Such lack of response
may be an artifact of experimental design such as
insufficient rooting volume in pots, which may restrict
the carbon storage capacity of roots (Berntson et al., 1993;
Conn et al., 1994). In addition, low nutrient availability may
decrease the capacity to accumulate additional carbon
under elevated CO2 conditions (Goudriaan and Ajtay, 1979;
Jarvis, 1989; Kramer, 1981; Oechel and Strain, 1985;
* Corresponding author. Tel.: +1 907 474 7652; fax: +1 907 474 6521.
E-mail address: ffjsc1@uaf.edu (J.S. Conn).
0167-8809/$ see front matter # 2005 Elsevier B.V. All rights reserved.
doi:10.1016/j.agee.2005.07.010
50
J.S. Conn, V.L. Cochran / Agriculture, Ecosystems and Environment 112 (2006) 4957
J.S. Conn, V.L. Cochran / Agriculture, Ecosystems and Environment 112 (2006) 4957
51
3. Results
3.1. Chamber performance
Average daily air and soil temperatures were not
significantly different (ANOVA, p > 0.05) between ANC
and OTC plots. The average daily air temperature measured
above the canopy for all plots between June 11 (when
52
J.S. Conn, V.L. Cochran / Agriculture, Ecosystems and Environment 112 (2006) 4957
Table 1
CO2 concentrations measured in ambient and elevated CO2 chambers
CO2 measurement
Mean of 08:0009:00 h and
16:0017:00 h measurements
(6/118/23)
Mean daily (7/258/23)
A + 175
A + 350
369 25
543 25
707 51
407 64
570 61
717 91
Table 2
Effects of CO2 treatments on potato gas exchange on July 13 and August 5 at Fairbanks, AK
Sampling date
Treatmenta
Photosynthesis
rate (mmol m2 s1)
Stomatal conductance
(m2 s1)
ANC
A + 350
22.1 a
22.5 a
0.31 b
0.12 a
130.2 a
321.3 b
August 5, 1994
ANC
A
A + 175
A + 350
17.4
15.4
19.7
23.6
0.49
0.34
0.24
0.26
249.6
242.7
368.2
492.3
ab
a
b
c
c
b
a
a
a
a
b
c
Means in a column within a date that are followed by different letters are significantly different (t-test, p < 0.05 for July 13; ANOVA, p < 0.05 and Duncans
MRT for August 5).
a
ANC = ambient CO2, no chamber; A = OTC, ambient CO2; A + 175 = OTC, ambient + 175 mmol mol1 CO2; A + 350 = OTC, ambient + 350 mmol mol1
CO2.
J.S. Conn, V.L. Cochran / Agriculture, Ecosystems and Environment 112 (2006) 4957
Table 3
First harvest ANOVA results
Variable
Treatment
r2
**
NS
NS
0.83
0.38
0.80
0.40
0.31
0.54
0.34
0.75
0.72
0.83
0.72
0.28
0.73
Block
Number of stems
Number of leaves (NL)
Number of flowers
Leaf area (LA)
Total leaf DW (LDW)
Specific leaf DW = LDW/LA
Average leaf DW = LDW/NL
Leaf % N
Total Stem DW (SDW)
Stem % N
Total flower DW (FDW)
Flower % N
Total shoot DW
(SHDW) = SDW + LDW + FDW
Number of tubers (NT)
**
NS
NS
NS
NS
NS
NS
NS
NS
NS
NS
**
NS
*
*
NS
NS
NS
NS
53
0.64
Total tuber WW
19:92 mmol mol1 CO2 above ambient 11685;
p < 0:001;
r 2 0:999:
Table 4
Effects of CO2 and chambers on potato at first harvest
Treatmenta
Leaf N
(%)
Stems
(number)
Flowers
(number)
Stem DW
(g)
Flower DW
(g)
Shoot DW
(g)
Stolon DW
(g)
ANC
A
A + 175
A + 350
5.35
5.54
5.11
4.92
2.5
2.8
2.3
2.0
64
95
65
49
311
396
252
247
33.1
45.4
27.7
17.7
1468
1727
1262
1328
15.3
18.7
17.2
12.4
ab
a
bc
c
ab
a
bc
c
b
a
b
b
b
a
c
c
ab
a
ab
b
ab
a
b
b
ab
a
a
b
Data are expressed on a per plant basis. Numbers in a column that are followed by different letters are significantly different as determined by Duncans multiple
range test ( p < 0.05).
a
ANC = ambient CO2, no chamber; A = OTC, ambient CO2; A + 175 = OTC, ambient + 175 mmol mol1 CO2; A + 350 = OTC, ambient +
350 mmol mol1 CO2.
Table 5
First harvest effects of CO2 on potato tuber gravity, root:shoot ratio, and biomass allocation
Treatmenta Tuber percent solids (%)Root:shoot ratioRoot allocation (%)Leaf allocation (%)Stem allocation (%)Flower allocation (%)Tuber allocation (%)
ANC
A
A + 175
A + 350
13.9
14.7
16.0
16.7
a
ab
bc
c
0.63
0.73
1.13
1.35
a
a
b
b
4.4
4.4
3.9
2.9
a
a
a
a
47.4
43.2
36.9
34.2
a
a
b
b
13.4 a
13.4 a
9.5 b
8.0 b
1.6
1.6
1.2
0.6
a
a
a
a
33.4
37.4
48.6
54.5
a
a
b
b
Numbers in a column that are followed by different letters are significantly different as determined by Duncans multiple range test ( p < 0.05).
a
ANC = ambient CO2, no chamber; A = OTC, ambient CO2; A + 175 = OTC, ambient + 175 mmol mol1 CO2; A + 350 = OTC, ambient +
350 mmol mol1 CO2.
54
J.S. Conn, V.L. Cochran / Agriculture, Ecosystems and Environment 112 (2006) 4957
Table 6
Second harvest ANOVA results
Variable
Block
Treatment
r2
Shoot DW
Shoot % N
Total tuber WW
Number 1 tuber WW
Knobby tuber WW
Cracked tuber WW
Tuber percent Solids
Total tuber DW
Tuber % N
Tuber sucrose
Tuber glucose
NS
NS
NS
NS
NS
NS
NS
NS
NS
NS
NS
NS
NS
NS
NS
NS
ND
NS
**
**
**
**
0.64
0.69
0.56
0.42
0.57
0.40
0.47
0.56
0.65
0.87
0.87
response to elevated CO2 at Fairbanks was that photosynthesis was limited by end product feedback inhibition (Guinn
and Mauney, 1980) induced by long days and low total
respiratory losses during short nights (<4.0 h). Extra starch
that accumulates in chloroplasts of leaves at high CO2
during the day is normally exported and used for dark
respiration at night (Ho, 1977; Kolbe and StephanBeckmann, 1997). In addition, since tuber initiation and
growth was not nearly as advanced on July 13 as on August
5, an adequate sink for starch transport may not have been
available. The longer night on August 5 (6.2 h) coupled with
a very strong sink in tubers for photosynthate could have
kept starch from accumulating in chloroplasts and downregulating photosynthesis. Studies by Stutte et al. (1996) add
support to this hypothesis. Norland potato net assimilation
rates were highest under a 12 h/12 h day/night photoperiod
and progressively decreased when plants were grown at 18 h
day/6 h night and 24 h light photoperiods. Moreover, leaf
starch levels increased when the dark period was decreased
to less than 12 h. Starch accumulation in potato leaves under
elevated CO2 was also observed by Goudriaan and de Ruiter
(1983), Sicher and Bunce (1999) and Schapendonk et al.
(2000).
An alternate explanation for the photosynthetic downregulation found in this study and in others is that the
response to elevated CO2 was limited by nitrogen supply.
Stitt and Krapp (1999) reviewed numerous experiments that
examined the interaction between N and response of plants
to elevated CO2 and found that photosynthetic downregulation was especially marked when nitrogen supply was
low. Adamsen et al. (2005) showed that when wheat
(Triticum aestivum L.) was grown at elevated CO2 as much
as 40% more mineral N was removed from the top 0.3 m of
soil than when grown at ambient CO2. At Fairbanks, while
first harvest foliar N was reduced 12.6% at the highest CO2
level (A + 350) compared to ambient CO2 chambers, there
were no significant effects of CO2 treatment on N
concentrations of shoots or tubers at the final harvest. In
the European CHIP experiment, potatoes grown at
680 mmol mol1 CO2 until harvest had significant reductions in nutrient concentrations both above-ground and in
tubers. Above-ground nitrogen declined 14.2%. Concentrations of nitrogen in tubers declined 6.4% (Fangmeier et al.,
Table 7
Effects of CO2 and chambers on potato at second harvest
Treatmenta
No. 1b tuber
wt. (kg ha1)
Tuber solids
(%)
Tuber sucrose
(mg g1)
Tuber glucose
(mg g1)
ANC
A
A + 175
A + 350
99.8
75.4
63.2
75.1
31431
18894
22264
25865
6380
3987
4787
5432
21800
12033
14398
20200
20.3
21.1
21.5
21.0
5.9
5.0
3.8
4.0
0.13
0.10
0.07
0.07
a
ab
b
b
a
b
b
b
Numbers in a column that are followed by different letters are significantly different as determined by Duncans multiple range test ( p < 0.05).
a
ANC = ambient CO2, no chamber; A = OTC, ambient CO2; A + 175 = OTC, ambient + 175 mmol mol1 CO2; A + 350 = OTC, ambient +
350 mmol mol1 CO2.
b
No. 1 tubers are >4.8 and <8.9 cm without defects.
J.S. Conn, V.L. Cochran / Agriculture, Ecosystems and Environment 112 (2006) 4957
55
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J.S. Conn, V.L. Cochran / Agriculture, Ecosystems and Environment 112 (2006) 4957
5. Conclusions
Potatoes grown at elevated CO2 at Fairbanks exhibited
enhanced tuber yields and changes in dry weight allocation
patterns. Biomass allocated to stems, leaves, and flowers
declined while that allocated to tubers increased with
elevated CO2. Root:shoot ratio was larger and percent leaf
nitrogen declined 12.6% with increased CO2. At the final
harvest, total tuber dry weight in the A + 350 chambers was
36% higher than in the A treatment. However, yields in
open-top chamber plots were lower than the no-chamber
plots possibly due to 714% less photosynthetic active
radiation in chambers. Whether global change will result in
higher yields of potatoes and other crops at high latitude
locations such as Fairbanks may depend as much on what
happens to cloud cover, light intensity, growing season
length and average temperature as it does on increasing
atmospheric CO2. This is because the current average
growing season temperature (15.8 8C) is at the lower end of
the range where increasing CO2 has been found to be
stimulatory to crop yields. Any increases in cloudiness that
reduce growing season temperature and PPFD could reduce
plant growth more than rising atmospheric CO2 levels can
stimulate growth. Finally, more field research is needed to
determine the interrelationships between N nutrition and
CO2 response in potato.
Acknowledgements
The authors thank Hugo Rogers and Lee Allen for help in
the design of the OTC system, for suggesting suppliers and
for providing other advice. Bill Saari built the equipment
building, the CO2 monitoring system, helped construct the
OTC system, and collected data. Richard Deck helped with
construction, collected data, and helped with data analysis.
Steve Prior made useful suggestions on an earlier version of
the manuscript.
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