Elevated carbon dioxide (CO2) influences photosynthesis (AN), transpiration (ET), and water use efficiency (WUE) for wellwatered potato (Solanum tuberosum L.). Little is known regarding effects of short-term drought and CO2 . Two experiments,
differing in the quantity of solar radiation, were conducted in soil-plant-atmosphere-research chambers. Plants were grown at
ambient (aCO2) or twice-ambient CO2 (eCO2) and received one of three irrigation treatments: no water stress (C), short-term
(1116 d) water-withholding during vegetative and post-tuber initiation stages (VR), or post-tuber initiation (R) only. Canopy
conductance to CO2 transfer () and water vapor (G v), light use efficiency (), daily AN, and ET decreased at the onset of each
drought and were correlated with volumetric water content. The rate of decrease was similar for R and VR. G v declined more
sharply than AN, resulting in higher WUE. Seasonal AN declined with the pattern of C > R > VR and was higher for eCO2 C
and R treatments. Seasonal WUE was higher for eCO2 at all irrigation treatments. Total dry matter, harvest index, and leaf area
were reduced (p < 0.05) for droughted treatments and total dry matter and harvest index were also higher for eCO2 VR pots.
Relative responses to drought and CO2 were similar among experiments, with greater magnitude of response under high solar radiation. Findings were similar to those reported under longer-term water-withholding studies, suggesting that interactions between CO2
and drought on carbon assimilation and water use are conserved across production zones with varying radiation and rainfall patterns.
2024
A g ro n o my J o u r n a l Vo l u m e 10 6 , I s s u e 6 2 014
Iersel and Kang, 2002). Using whole canopy gas exchange measurements, based on averaged CO2 and water vapor exchange
rates across multiple plants in the same enclosure, Fleisher et
al. (2012; 2008a, 2008b) observed higher canopy net and gross
photosynthesis (AN and AG), reduced transpiration (ET), higher
radiation use efficiency and WUE, yet similar leaf area production, for eCO2 vs. aCO2 grown plants
Potato drought responses are influenced by the duration and
timing of the water with-holding event with respect to developmental stage (Dalla Costa et al., 1997; Iqbal et al., 1999;
Kang et al., 2004; Kashyap and Panda, 2003; Schafleitner,
2009). Few articles in the literature report the impact of eCO2
on potato production and drought; however, under aCO2
responses include reduced leaf area, canopy size and duration
(Jefferies, 1995; Fleisher et al., 2008a, 2008b; Lahlou et al.,
2003; van Loon, 1981), decreased dry matter production and
yield (Dalla Costa et al., 1997; Deblonde and Ledent, 2001;
Fleisher et al., 2008a, 2008b; Schittenhelm et al., 2006; van
Loon, 1981), with higher harvest index and WUE (Belanger et
al., 2001; Fleisher et al., 2008a; Trebejo and Midmore, 1990).
Under aCO2 in enclosure studies, Vos and Groenwold (1989)
measured a larger effect of drought on canopy stomatal conductance than AN, resulting in improved whole plant WUE over a
period of 33 d. Dalla Costa et al. (1997) found that AN of plant
canopies that were water-stressed during vegetative growth was
more sensitive, and less able to recover to re-watering events,
as compared to plants water-stressed following tuber maturity.
Only two studies on potato, eCO2 , and drought interactions
were reported (Fleisher et al., 2008a, 2008b), in which the
authors found droughted plants had similar total biomass, but
higher harvest index, radiation use efficiency, and WUE compared with aCO2 grown potato. It was unclear whether these
responses to long-term drought would manifest under the more
likely to occur short-term water with-holding periods.
Basu et al. (1999) indicated that tuber sink strength could
mediate drought-based photosynthetic feedback inhibition
under aCO2 . In addition to reduced stomatal conductance,
which has been identified as the primary mechanism for
improving WUE in C3 (Finnan et al., 2005) and C4 crops
in particular (Allen et al., 2011), growth at eCO2 results in a
larger plant carbon assimilate pool which has the potential to
form an even stronger tuber sink, potentially increasing the
capacity for reducing photosynthetic feedback inhibition as
compared to aCO2 . Since this drought tolerance mechanism
depends on the timing of tuber initiation and bulking stages,
interactions with periodic water-deficit, CO2 levels, and phenology need to be investigated. Research has also shown that
irradiance level influences the degree to which CO2 enrichment enhances leaf gas exchange in potato with a proportionately larger increase in these responses observed at lower light
levels (Ku et al., 1977; Wheeler et al., 1991). Based on these
observations, we speculate that eCO2 would reduce effects of
short-term drought on potato gas exchange and agronomic
response, that this moderating effect would be more pronounced during drought occurring post tuber-initiation stage
than at a prior vegetative event, and the relative degree of this
reduction would be greater under lower irradiance levels.
Sunlit enclosures were used to study effects of CO2 and shortterm droughts, when applied at different developmental stages,
2025
Table 1. Timing of drought cycles in days after emergence (DAE), average daily light integral (PAR), and averaged day (Tday) and night (Tnight) air temperatures, and 24 h relative humidity (RH) across all treatments for experiments E1 and E2.
Event
Drought 1
Drought 2
Season
Dates
DAE
1127
4559
171
PAR
mol m2 d1
41.7 10.3
45.5 8.7
43.9 10.1
E1
Tday
Tnight
C
22.0 0.17 17.0 0.16
21.9 0.47 17.5 0.61
22.0 0.21 17.3 0.36
RH
%
76.7 0.52
69.9 3.89
73.4 2.35
Dates
DAE
1329
4152
174
E2
PAR
Tday
Tnight
RH
mol m2 d1 C
%
29.3 15.0 22.1 0.12 17.1 0.04 78.9 0.06
15.7 5.9
22.0 0.17 17.0 0.18 79.9 0.83
24.7 12.4 22.0 0.08 17.1 0.10 79.0 0.11
[1]
AG =
a I tC
aI +tC
[2]
eC (TC ) - ea
lET = Gv
pa
[3]
ec(Tc) and ea are the vapor pressures at the canopy surface and
in the air, and pa is atmospheric pressure.
Statistical Analysis
To test for differences due to solar radiation levels between
experiments, dry matter data were initially modeled as a twofactor factorial design with experiment as a random block. The
likelihood ratio statistic (Littell et al., 1996) was used to determine if this block effect was significant for each dry matter
response. The SAS MIXED procedure (SAS Institute, 2009)
was used, with chamber as a random effect, for analysis of variance of dry matter (n = 6) and time-averaged gas exchange data
(n 11, depending on experiment and drought cycle as in Table
1) during each drought. Mean separations were conducted with
Fishers protected LSD ( = 0.05). For dry matter analyses,
individual pots were used as pseudo-replicates as per Fleisher et
al. (2009) and Reddy and Zhao (2005). Covariance analyses,
using average PAR during the given harvest stage, were also
used to test for commonality of responses to CO2 and H2O
treatment factors between the two experiments for dry matter
responses. Linear regression and contrasts were conducted
using SAS Proc MIXED to compare differences in all other gas
Fig. 1. Average daily volumetric water contents (VWC, n = 7) and mid-day leaf water potentials (LWP, n = 3) vs. day after emergence during two
drought cycles for experiments (a, b) E1 and (c, d) E2 at (a, c) ambient CO2 (aCO2) and (b, d) elevated CO2 (eCO2). H2O symbols: circle-C (control),
inverted triangle-R (drought applied at post tuber-initiation stage), square-VR (drought applied at vegetative and post tuber-initiation stages). Open
symbols for VWC and closed symbols for LWP. Standard errors indicated.
2027
Table 2. Decline in gas exchange parameters (canopy light use efficiency, canopy conductance to carbon dioxide transfer, Gv canopy conductance to water vapor), normalized with respect to ambient (aCO2) or elevated (eCO2) CO2 control chamber, vs. decreasing volumetric water content
(VWC) during drought cycles (DC) in experiment E1 and E2 for H2O treatments VR (droughted at vegetative and post tuber-initiation stages) and R
(droughted at post tuber-initiation stage). Slopes and standard error of the estimate are shown. Letters indicate significant differences within a given
CO2 treatment. Slopes that were significantly different between CO2 treatments indicated by * ( = 0.5) or ( = 0.1).
Experiment
DC
E1
1
2
E2
1
2
* Significant at = 0.05.
Significant at = 0.1.
H2O
treatment
VR
VR
R
VR
VR
R
aCO2
eCO2
aCO2
eCO2
aCO2
eCO2
%1
0.024 (0.005)* 0.045 (0.003)*
0.03 (0.008)*
0.055 (0.005)*
0.037 (0.005)*
0.058 (0.01)*
0.031 (0.007) 0.020 (0.003)
0.052 (0.009)*
0.030 (0.006)a*
0.021 (0.002)*
0.043 (0.007)*
0.035 (0.007)
0.022 (0.004)
0.047 (0.005)*
0.079 (0.006)b*
0.023 (0.002)*
0.056 (0.007)*
0.027 (0.006)
0.026 (0.004)
0.056 (0.02)
0.042 (0.005)
0.037 (0.005)
0.036 (0.003)
0.03 (0.008)
0.02 (0.002)
0.083 (0.03)
0.094 (0.02)a
0.047 (0.009)a
0.043 (0.003)
0.023 (0.005)
0.022 (0.002)
0.045 (0.013)
0.027 (0.004)b
0.026 (0.004)b* 0.044 (0.007)*
2028
Gv
Table 3. Averaged gas exchange values (ANcanopy net photosynthesis; ETcanopy evapotranspiration rate; WUEcanopy water use efficiency;
canopy light use efficiency; canopy conductance to carbon dioxide transfer; Gv canopy conductance to water vapor) for drought cycles (DC) 1 and
2 for E1 (n = 15 and 13, respectively) and E2 (n = 15 and 10, respectively). H2O symbols: Cnon-water stressed control; Rtreatments droughted at
post tuber-initiation stage only; VRtreatments droughted at post tuber-initiation and vegetative growth. Least square differences (LSD) within each
CO2 grouping are provided. Letters indicate significant differences within a given CO2 treatment ( = 0.05) and * indicates differences across CO2
treatments for a given H2O grouping ( = 0.05).
Experiment DC
E1
H2O
treatment
E2
C
VR
LSD
C
R
VR
LSD
C
VR
LSD
C
R
VR
LSD
AN
aCO2
ET
eCO2
aCO2
WUE
eCO2
mol m2 d1
0.85a
0.85a
319a* 261a*
0.49b
0.46b
167b* 104b*
0.14
0.19
25
32
1.25a* 1.58a*
453a* 336a*
0.52b
0.73b
107b
81b
0.53b
0.66b
101b
69b
0.16
0.16
28
25
0.55a
0.76
480a* 410a*
0.27b* 0.54*
125b
106b
0.14
0.25
55
40
0.38
0.48a
312a* 335a*
0.29
0.20b
117b
118b
0.36
0.33a
121b
104b
0.14
0.15
33
28
aCO2
eCO2
mmol mol1
2.7*
3.6a*
2.9*
5.7b*
0.6
1.4
2.5a*
5.1a*
6.0b*
9.2b*
5.2b*
9.6b*
1.0
1.5
1.1a*
1.9a*
4.6b
7.2b
2.3
1.7
1.4a*
2.0a*
4.2b
3.5a
3.5b
6.4b
1.0
1.5
aCO2
eCO2
aCO2
Gv
eCO2
mm s1
4.29a* 2.84a*
2.21b* 1.12b*
0.50
0.60
5.41a* 4.45a*
2.42b
1.75b
2.22b
1.52b
0.64
0.47
4.43a
3.43a
2.58b
2.04b
1.3
0.90
7.55a* 5.01a*
4.85b* 2.45b*
5.33b* 2.56b*
1.06
0.63
aCO2
eCO2
mmol m2 s1
679a*
460a*
356b*
203b*
86
72
525a
471a
120b
125b
147b
121b
54
49
1203a* 717a*
300b
257b
234
109
1079a* 680a*
235b
305b
337b
231b
111
74
* Significant at = 0.05.
than AN. For example, averaged across CO2 and H2O, G v was
reduced 51 to 74% as a result of the droughts while declined
40 to 60%. Light use efficiency was negatively impacted by the
droughts, particularly in E1. In nearly all cases there was no
evidence to suggest that the gas exchange responses of twicedroughted VR pots were more or less sensitive to a second
drought post tuber-initiation period than the single-droughted
R pots. Thus, previously droughted plants did not exhibit any
additional drought tolerance mechanisms to the additional
drought cycle, at least when assessed at the canopy level during
the drought itself.
Seasonal Gas Exchange and
Dry Matter Production
These transient gas exchange responses were reflected in the
whole season responses (Table 4). Cumulative AN decreased
with increasing drought frequency with the pattern of C >
R > VR, although such differences were more substantial in E1
than E2. Similar to responses during the water withholding
periods, seasonal ET of droughted pots generally declined to
a greater extent than AN as compared to the C values, resulting in higher WUE over the course of the season (Table 4).
Some variation existed with VR WUE responses, likely due
to higher soil evaporation as a result of sparser canopy growth.
Treatments with eCO2 showed consistently higher AN than
aCO2 across both experiments with either lower, or similar E
values, thus resulting in higher WUE regardless of H2O treatment. Less than 14% error between end-of-season dry matter
and cumulative AN was observed in all treatments (Table 4,
Error column). These C balance results are typical in canopy
gas exchange studies (Van Iersel and Kang, 2002) and indicate
the observations of seasonal AN were compatible with total dry
matter production. Slopes from the regression of cumulative
AN on time served as an analogue for plant growth rate (Fig. 4),
and confirmed that decreased AN in the VR and R treatments
was due to the drought cycles (Table 5). For example, growth
2029
Table 4. End-of-season (cumulative) canopy net photosynthesis (AN), evapotranspiration (ET), water use efficiency (WUE), dry matter, and percent
difference (error) between end-of-season dry matter production and AN. H2O symbols as defined in Table 3.
Experiment E1
CO2
aCO2
eCO2
Experiment E2
ET
WUE
Dry mass
Error
ET
WUE
Dry mass
mol CO2 m2
mol H2O m2
mmol mol1
mol CO2 m2
mol CO2 m2
mol H2O m2
mmol mol1
mol CO2 m2
64.9
24,348
2.67
75.4
13.9
41.4
20,971
1.97
40.4
2.6
R
VR
54.4
44.7
18,284
15,339
2.98
2.91
63.2
45.5
13.9
1.9
40.5
35.9
18,031
12,259
2.25
2.93
38.4
38.6
5.3
6.9
C
R
VR
90.4
77.5
40.9
20,580
16,825
11,762
4.39
4.61
3.48
92.2
82.1
46.4
2.1
5.6
11.8
53.6
57.6
35.0
22,028
18,457
11,019
2.43
3.12
3.18
55.4
52.8
39.6
3.2
9.0
11.7
H2O
AN
AN
Error
%
Includes all dry matter from partial and final harvests. Dry matter was converted to mol CO2 basis based on average tissue carbon content of 40% for purposes of C
balance comparison.
Table 5. The rate of increase in canopy AN (mol CO2 m 2 d 1) during drought cycles 1 and 2 as obtained from Fig. 3 (r 2 = 0.95 or higher for all regressions). H2O symbols as defined in Table 3. Standard errors of the estimate indicated in parentheses. Letters indicate significant differences ( = 0.05)
within a given CO2 treatment and * indicates differences across CO2 treatments for a given H2O grouping ( = 0.05).
Experiment E1
Cycle
Experiment E2
H 2O
aCO2
eCO2
aCO2
eCO2
Drought 1
C
VR
0.86 (0.04)a
0.58 (0.02)b*
0.79 (0.07)a
0.44 (0.03)b*
0.72 (0.07)a*
0.45 (0.03)b*
1.01 (0.09)a*
0.64 (0.04)b*
Drought 2
C
R
VR
1.19 (0.08)a*
0.59 (0.05)b*
0.69 (0.04)b*
1.78 (0.09)a*
0.71 (0.06)c*
0.88 (0.04)b*
0.58 (0.05)a*
0.60 (0.03)a
0.62 (0.02)a*
0.85 (0.07)a*
0.55 (0.04)b
0.54 (0.03)b*
* Significant at = 0.05.
Fig. 4. Cumulative canopy net assimilation (AN) for all H2O treatments vs. days after emergence for experiments (a, b) E1 and (c, d) E2 at (a, c) ambient
CO2 (aCO2) and (b, d) elevated CO2 (eCO2). Drought cycle timing and durations are indicated with shaded lines.
2030
Table 6. Total and leaf dry matter, leaf area, and harvest index (HI) at final harvest for experiments E1 and E2 expressed on a per pot basis (n = 6).
Analysis of variance and least square differences (LSD) indicated. Symbols as defined in Table 3.
Experiment E1
CO2
H2O
aCO2
C
R
VR
LSD
C
R
VR
eCO2
LSD
CO2
H2O
CO2 H2O
Total
Leaf
g pot1
232a*
46.7
167b
46.2
126c
42.3*
28.3
19.4
273a*
194b
136c
42
*
***
63a
47a
27b*
11.7
ns
**
*
Area
Experiment E2
HI
Total
cm2 pot1
10,123a
7,163b
8,353a*
1,574
0.43a
0.39b
0.32b*
0.08
12,647a
8,063b
6,029b*
2,309
ns
***
*
0.47b
0.47b
0.60a*
0.13
**
ns
*
Leaf
g pot1
122*
33.1a
96*
22.1b*
101
29.4a*
25.5
7.3
143a*
152a*
105b
46.2
*
ns
34.2b
46.9a*
19.4c*
8.8
*
*
***
Area
HI
cm2 pot1
9,857a
6,417b*
9,127a*
1,429
0.20a
0.39b
0.51c
0.19
10,277a
10,666a*
4,599b*
2,822
ns
**
***
0.43a
0.50ab
0.61b
0.11
**
**
ns
* Significant at 5%.
** Significant at 1%.
*** Significant at 0.1%.
ns, nonsignificant.
Significant at 10%.
for other H2O treatments at either CO2 level each day of the
respective experiments. Comparison on a given day between
experiments is strongly influenced by differences in canopy
development (a result of the radiation differences in E1 and
E2), however leaf area, while generally smaller in E2 than E1,
was not proportionately reduced in response to the drought
treatments to the same extent as AN, total or leaf dry matter
production (Table 6). This was largely a result of differences in
specific leaf area for the plant canopies which averaged 17.9
1.63 m2 kg1 in E1 and 27.9 3.35 m2 kg1 in E2, indicating
leaves were substantially thinner in E2 as a result of acclimation
to the lower light environment. It should be further emphasized that such values are instantaneous rates. When photosynthetic data are integrated over the course of the day or season
(such as the data in Tables 3 and 4), substantial differences were
observed between E1 and E2 due to reduced daily and seasonal
radiation levels (Table 1). Changes in C allocation and leaf
thickness as a result of the different radiative environments
between E1 and E2 impacted the degree to which drought
results were manifested, particularly in terms of the extent to
which dry matter and seasonal AN were reduced with respect
to the control treatments (Table 4).
Although the per-pot dry matter data was evaluated separately in each experiment (Table 6), an analysis of covariance
was conducted separately using the pooled data set across E1
and E2, with mean daily PAR during each H2O treatment
period (Table 1) used as the concomitant variable. With the
exception of harvest index, PAR was found to have a significant
influence on all dry matter and leaf area responses (data not
shown). Analyzed in this way, total dry matter and tuber production were significantly impacted by H2O and CO2, while leaf
and leaf area production showed by H2O and interactive effects.
DISCUSSION
The effects of the short-term water-withholding cycles on dry
matter production, HI, AN, ET, and WUE at the two different
CO2 levels were analogous to those reported from a long-term
water-withholding study (Fleisher et al., 2008a). In that study,
2031
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