With changing global precipitation patterns, will the growth of prairie plants be more constrained...
1
With changing global precipitation patterns, will the growth of prairie plants be more constrained by reduced rainfall frequency or reduced rainfall volume? Adam Schneider*, Molly Kreiser, Greg Nelson ● Faculty Mentor: Tali Lee Department of Biology-University of Wisconsin Eau Claire *[email protected] Water deficits are a common limiting factor of plant growth. In North America, rainfall gradients are a key determinant of ecosystem type and primary productivity, particularly among grasslands (Fay et. al., 2003). Rainfall patterns are one of a number of climate factors that are expected to change substantially in the coming decades. Predictions for the Upper Midwest show only a slight increase in total rainfall, but a dramatic change in the distribution of rainfall, including severe June-August droughts, and larger spring rain events (Winter & Eltahir, 2012; Hayhoe et. al., 2009). Many studies have looked at the effects of drought on plants, but few have independently compared the two ways a plant might experience reduced water availability: fewer rain events, or less water per event. A decade-long field study of a tallgrass prairie in Kansas found that for some species, reducing the frequency of rainfall events while keeping total water volume constant reduced soil water availability as much as reducing the rainfall amount. Also, grass species were found to compensate for changing water availability through physiological adjustments (Fay et. al., 2003). However, the way these findings relate to other community types, species, and functional groups is uncertain. And, while field studies allow more natural conditions, growth in controlled conditions can provide additional information, such as access to below-ground plant parts. In this greenhouse experiment we investigated responses of the legume Lupinus perennis and the grass Agropyron repens to 50% reductions in rainfall frequency, total rainfall volume, or both (Table 1). Our objective was to contrast the effects of reduced rainfall amount with a more extreme distribution of rainfall on plant by measuring leaf senescence, biomass accumulation, photosynthetic and stomatal conductance rates, and for , the amount of nitrogen (N) derived from symbiotic N fixation. Introduction Materials and Methods Experiment Design • Plants grown in cylindrical pots (5.6 cm dia., X 30 cm; 3.0 L) • Soil collected from experimental grasslands at Cedar Creek Ecosystem Science Reserve in east-central Minnesota. • Single individuals of Agropyron repens and Lupinus perennis grown under standard greenhouse conditions for fourteen weeks; 14 hour photoperiod. • Water availability was manipulated by reducing the total water volume by 50%, reducing the frequency of watering events by 50% or both (N=88, Table 1). o Plants were well watered until they developed two true leaves, at which point they were thinned to one per pot and treatments began. o Watering treatments were determined from a pilot study to simulate rain volumes, frequencies and resulting soil moisture availabilities experienced at Cedar Creek Ecosystem Science Reserve. • Each pot was fertilized 1.70 mg of NH 4 NO 3 enriched with 15 N (0.3850 atom% 15 N) four times to determine the proportion of N derived from N 2 fixation in Lupinus by isotope analysis. Measurements • Volumetric % soil moisture (immediately before and one day after each watering event) • Nitrogen stable isotope ratios (atom % 15 N vs 14 N) • Total number of leaves per plant that were expanding, expanded, senescing, or fully senesced (during weeks 5, 7, and 13) • Total accumulated shoot and root biomass • Leaf gas exchange (net photosynthetic CO 2 uptake and stomatal conductance of H O) using a portable infrared The watering treatments strongly affected the soil water content of each pot though remained realistic to what a native Upper-Midwest prairie plant may experience during a growing season in terms of both water availability and rainfall frequency (Table 1, Fig. 1). Specific responses varied by species, but in general, reduced rainfall volume was found to be more limiting than reduced rainfall frequency. It is well established that N fixation by legumes is sensitive to drought, though research has been mostly limited to agricultural species (Serraj et. al. 1999). In our study, reducing watering frequency did not affect N fixation, whereas reducing in total water volume significantly reduced the proportion of plant N derived from fixation in Lupinus. Moreover, imposing both forms of reduced water availability resulted in a synergistic decrease in N fixation (Fig. 2). This suggests that it is only the reduction in rainfall—not necessarily the frequency of rainfall events—that is mostly responsible for inhibiting legume N fixation. Similar responses were found when leaf senescence and biomass were measured. Lupinus showed an increase in the proportion of senesced leaves and a reduction in total dry biomass with decreased water availability, but these effects were similarly only significant in cases where the total volume of water received was reduced (Fig. 3B, 4). The proportion of Agropyron leaves senescing at a given time and total dry biomass were not significantly different from the control in any of the treatments, suggesting that this species is better adapted to drought (Fig. 3B, 4). Watering treatments reduced leaf net photosynthetic rates in Lupinus only. However, due to a greater reduction in stomatal conductance (water loss) than Lupinus, Agropyron increased its water use efficiency for reduced volume and reduced volume and frequency treatments compared to the control (Fig. 5). This is consistent with the prevailing generality that grasses are better adapted to drought than forbs (Fay et. al. 2003). For every growth parameter studied, a 50% reduction in watering frequency did not significantly affect Discussion and Conclusions Acknowledgement s Funding sources: • UW-Eau Claire Differential Tuition • UW-Eau Claire Office of Research and Sponsored Programs • Blugold Fellowship (A. Schneider) Also, special thanks to Susan Barrott and Cedar Creek Ecosystem Science Reserve for soil moisture data, stable isotope analysis, and soil for the experiment. Results ↓Frequency ↓Volume ↓Vol. / ↓Freq. 0 10 20 30 40 Watering Treatment % Change in proportion of leaves in senescence ** ** Figure 3. (A) Reduced water treatments did not affect the number of leaves produced by Agropyron or Lupinus. However, (B) 50% reduced rainfall volume increased the proportion of leaves in senescence of Lupinus, although Agropyron senescence was not significantly affected. Leaves were counted at weeks 5, 7, and 13. Error bars indicate standard error; asterisks indicate a significant difference from the control (p<0.05). Control ↓Frequency ↓Volume ↓Vol. / ↓Freq. 0 5 10 15 20 25 30 Agropyron Lupinus Watering Treatment Total Number of Leaves Produced A B Effects on Leaf Production and Senescence Figure 2. Compared to the control, Lupinus showed a reduction in the proportion of N derived from fixation of 13% in a reduced volume situation and a 22% reduction in a reduced frequency and volume situation. Treatments not connected by the same letter are significantly different (p<0.05). Control ↓ Frequency ↓ Volume ↓ Freq. / ↓ Vol. 0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1 Watering Treatment Percent N derived from Fixation B A C A B A C A Effects on Legume Nitrogen Fixation References Fay, P.A., Carlisle, J.D., Knapp, A.K., Blair, J.M, Collins, S.L., 2003. Productivity responses to altered rainfall patterns in a C 4 -dominated grassland. Oecologia, 137(2), 245- 251. Hayhoe, K., VanDorn, J., Naik, V., Wuebbles, D., 2009. Climate Change in the Midwest: Projections of Future Temperature and Precipitation. Cambridge, MA: Union of Concerned Scientists. Serraj. R., Sinclair, T.R., Purcell, L.C., 1999. Symbiotic N 2 fixation response to drought. Journal of Experimental Botany 50:143-155 Winter, J.M., Eltahir, E.A.B., 2012. Modeling the Effects on Soil Moisture 0 11 0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0 18.0 20.0 22.0 24.0 Control Reduced Frequency Reduced Volume Reduced Freq. and Vol. Mean Days Since Last Reduced-Frequency Watering Volumetric Soil Moisture (%) 1 6 7 0 11 0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0 18.0 20.0 22.0 24.0 Mean Days Since Last Reduced-Frequency Watering Volumetric Soil Moisture (%) 1 6 7 Figure 1. Effect of watering treatments on soil moisture of pots containing single individuals of Agropyron repens and Lupinus perennis. Soil moisture was measured in all pots immediately before and one day after any watering event. Points represent the mean ± SE of eleven replicates over eight watering cycles (n=88). Orange bars indicate mean soil moisture field data at six and twelve days after a >1 cm rain event in experimental grasslands at Cedar Creek Ecosystem Science Reserve in east-central Minnesota (May- July, 2000-2010). Agropyron Lupinu s ↓ Frequency ↓ Volume ↓ Freq. / ↓ Vol. Control 0.0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 Root to Shoot Ratio Control ↓ Freq. ↓ Vol. ↓ Freq. / ↓ ... 0.0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0 9.0 Total Dry Biomass (g) AB Control ↓ Freq. ↓ Vol. ↓ Freq./ ↓ Vol 0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0 9.0 Watering Treatment Watering Treatment Figure 4. Biomass accumulation was significantly reduced in Lupinus but not in Agropyron under decreased water availability. The ratio of accumulated root to shoot biomass did not change in Lupinus, but there appeared to be a weak antagonistic effect between decreased volume and decreased frequency in Agropyron. Treatments not connected by the same letter are significantly different (p<0.05). Bars represent the mean ± SE of eleven replicates. Agropyron Lupinus Effects on Root, Shoot, and Total Biomass AB A B B C A A B AB AB A Effects on Leaf Physiology and Water Use Efficiency Watering Treatment Watering Treatment C Figure 5. Percent changes in leaf physiology of plants grown with reduced water availability compared to the control. Effect on leaf net photosynthetic rates was variable (A); however, due to a decrease in conductance (B), the water use efficiency (CO 2 assimilation per unit water loss) increased significantly in Agropyron under combined water stress conditions whereas Lupinus had a more variable response (C). Error bars represent standard error; asterisks indicate a significant difference between treatments and the control (** = p<0.05; * = p<0.10). -90 -80 -70 -60 -50 -40 -30 -20 -10 0 10 % Change in Conductance -20 0 20 40 60 80 100 % Change in Water Use Efficiency -70 -60 -50 -40 -30 -20 -10 0 10 20 30 Agropyron Lupinus % Change in Photosynthesis ** ** A B C Watering Treatment Watering Treatment * * * * * Water Treatments Table 1. Volume and frequency of water given to plants in each of four Average days between watering Volume per event (mL) Control 5.7 300 ↓Frequenc y 11.4 600 ↓Volume 5.7 150 ↓Frequenc y / ↓Volume 11.4 300 Before watering After watering Measuring soil moisture
-
Upload
griffin-austin -
Category
Documents
-
view
216 -
download
0
Transcript of With changing global precipitation patterns, will the growth of prairie plants be more constrained...
With changing global precipitation patterns, will the growth of prairie plants be more constrained by reduced rainfall frequency or reduced rainfall volume?
Adam Schneider*, Molly Kreiser, Greg Nelson ● Faculty Mentor: Tali Lee Department of Biology-University of Wisconsin Eau Claire
Water deficits are a common limiting factor of plant growth. In North America, rainfall gradients are a key determinant of ecosystem type and primary productivity, particularly among grasslands (Fay et. al., 2003). Rainfall patterns are one of a number of climate factors that are expected to change substantially in the coming decades. Predictions for the Upper Midwest show only a slight increase in total rainfall, but a dramatic change in the distribution of rainfall, including severe June-August droughts, and larger spring rain events (Winter & Eltahir, 2012; Hayhoe et. al., 2009).
Many studies have looked at the effects of drought on plants, but few have independently compared the two ways a plant might experience reduced water availability: fewer rain events, or less water per event. A decade-long field study of a tallgrass prairie in Kansas found that for some species, reducing the frequency of rainfall events while keeping total water volume constant reduced soil water availability as much as reducing the rainfall amount. Also, grass species were found to compensate for changing water availability through physiological adjustments (Fay et. al., 2003). However, the way these findings relate to other community types, species, and functional groups is uncertain. And, while field studies allow more natural conditions, growth in controlled conditions can provide additional information, such as access to below-ground plant parts.
In this greenhouse experiment we investigated responses of the legume Lupinus perennis and the grass Agropyron repens to 50% reductions in rainfall frequency, total rainfall volume, or both (Table 1). Our objective was to contrast the effects of reduced rainfall amount with a more extreme distribution of rainfall on plant function by measuring leaf senescence, biomass accumulation, photosynthetic and stomatal conductance rates, and for Lupinus, the amount of nitrogen (N) derived from symbiotic N2 fixation.
Introduction
Materials and MethodsExperiment Design• Plants grown in cylindrical pots (5.6 cm dia., X 30 cm; 3.0 L)• Soil collected from experimental grasslands at Cedar Creek Ecosystem
Science Reserve in east-central Minnesota.• Single individuals of Agropyron repens and Lupinus perennis grown
under standard greenhouse conditions for fourteen weeks; 14 hour photoperiod.
• Water availability was manipulated by reducing the total water volume by 50%, reducing the frequency of watering events by 50% or both (N=88, Table 1). o Plants were well watered until they developed two true leaves, at
which point they were thinned to one per pot and treatments began.o Watering treatments were determined from a pilot study to simulate
rain volumes, frequencies and resulting soil moisture availabilities experienced at Cedar Creek Ecosystem Science Reserve.
• Each pot was fertilized 1.70 mg of NH4NO3 enriched with 15N (0.3850 atom% 15N) four times to determine the proportion of N derived from N2 fixation in Lupinus by isotope analysis.
Measurements• Volumetric % soil moisture (immediately before and one day after each
watering event) • Nitrogen stable isotope ratios (atom % 15N vs 14N)• Total number of leaves per plant that were expanding, expanded,
senescing, or fully senesced (during weeks 5, 7, and 13)• Total accumulated shoot and root biomass• Leaf gas exchange (net photosynthetic CO2 uptake and stomatal
conductance of H2O) using a portable infrared gas exchange analyzer between watering treatments (week 11)
Statistical Analyses• Data were analyzed using ANOVA (JMP 8.0.2.2).
The watering treatments strongly affected the soil water content of each pot though remained realistic to what a native Upper-Midwest prairie plant may experience during a growing season in terms of both water availability and rainfall frequency (Table 1, Fig. 1).
Specific responses varied by species, but in general, reduced rainfall volume was found to be more limiting than reduced rainfall frequency. It is well established that N fixation by legumes is sensitive to drought, though research has been mostly limited to agricultural species (Serraj et. al. 1999). In our study, reducing watering frequency did not affect N fixation, whereas reducing in total water volume significantly reduced the proportion of plant N derived from fixation in Lupinus. Moreover, imposing both forms of reduced water availability resulted in a synergistic decrease in N fixation (Fig. 2). This suggests that it is only the reduction in rainfall—not necessarily the frequency of rainfall events—that is mostly responsible for inhibiting legume N fixation.
Similar responses were found when leaf senescence and biomass were measured. Lupinus showed an increase in the proportion of senesced leaves and a reduction in total dry biomass with decreased water availability, but these effects were similarly only significant in cases where the total volume of water received was reduced (Fig. 3B, 4). The proportion of Agropyron leaves senescing at a given time and total dry biomass were not significantly different from the control in any of the treatments, suggesting that this species is better adapted to drought (Fig. 3B, 4).
Watering treatments reduced leaf net photosynthetic rates in Lupinus only. However, due to a greater reduction in stomatal conductance (water loss) than Lupinus, Agropyron increased its water use efficiency for reduced volume and reduced volume and frequency treatments compared to the control (Fig. 5). This is consistent with the prevailing generality that grasses are better adapted to drought than forbs (Fay et. al. 2003).
For every growth parameter studied, a 50% reduction in watering frequency did not significantly affect the plant as long as there no overall reduction in total water volume. However, if volume was reduced by 50%, Lupinus showed a reduction in growth and nitrogen fixation and Agropyron showed changes in leaf physiology (water use efficiency) to compensate. Reductions in volume and frequency together showed additive or synergistic effects. In the future, predictions of vegetation responses to climate change could be improved by considering these two dimensions of water availability independently.
Discussion and Conclusions
AcknowledgementsFunding sources:
• UW-Eau Claire Differential Tuition• UW-Eau Claire Office of Research and
Sponsored Programs• Blugold Fellowship (A. Schneider)
Also, special thanks to Susan Barrott and Cedar Creek Ecosystem Science Reserve for soil moisture data, stable isotope analysis, and soil for the experiment.
Results
↓Frequency ↓Volume ↓Vol. / ↓Freq.0
10
20
30
40
Watering Treatment
% C
hang
e in
pro
porti
on
of le
aves
in se
nesc
ence
**
**
Figure 3. (A) Reduced water treatments did not affect the number of leaves produced by Agropyron or Lupinus. However, (B) 50% reduced rainfall volume increased the proportion of leaves in senescence of Lupinus, although Agropyron senescence was not significantly affected. Leaves were counted at weeks 5, 7, and 13. Error bars indicate standard error; asterisks indicate a significant difference from the control (p<0.05).
Control ↓Frequency ↓Volume ↓Vol. / ↓Freq.0
5
10
15
20
25
30 Agropyron Lupinus
Watering Treatment
Tota
l Num
ber o
f Lea
ves P
rodu
ced
A B
Effects on Leaf Production and Senescence
Figure 2. Compared to the control, Lupinus showed a reduction in the proportion of N derived from fixation of 13% in a reduced volume situation and a 22% reduction in a reduced frequency and volume situation. Treatments not connected by the same letter are significantly different (p<0.05).
Control ↓ Frequency ↓ Volume ↓ Freq. / ↓ Vol.0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1
Watering Treatment
Perc
ent N
der
ived
from
Fix
ation
B
A
C
A
B
A
C
A
Effects on Legume Nitrogen Fixation
ReferencesFay, P.A., Carlisle, J.D., Knapp, A.K., Blair,
J.M, Collins, S.L., 2003. Productivity responses to altered rainfall patterns in a C4-dominated grassland. Oecologia, 137(2), 245-251.
Hayhoe, K., VanDorn, J., Naik, V., Wuebbles, D., 2009. Climate Change in the Midwest: Projections of Future Temperature and Precipitation. Cambridge, MA: Union of Concerned Scientists.
Serraj. R., Sinclair, T.R., Purcell, L.C., 1999. Symbiotic N2 fixation response to drought. Journal of Experimental Botany 50:143-155
Winter, J.M., Eltahir, E.A.B., 2012. Modeling the hydroclimatology of the midwestern United States. Part 2: future climate. Climate Dynamics, 38:595-61.
Effects on Soil Moisture
0 110.0
2.0
4.0
6.0
8.0
10.0
12.0
14.0
16.0
18.0
20.0
22.0
24.0Control
Reduced Frequency
Reduced Volume
Reduced Freq. and Vol.
Mean Days Since Last Reduced-Frequency Watering
Volu
met
ric S
oil M
oist
ure
(%)
1 6 7
0 110.0
2.0
4.0
6.0
8.0
10.0
12.0
14.0
16.0
18.0
20.0
22.0
24.0
Mean Days Since Last Reduced-Frequency Watering
Volu
met
ric S
oil M
oist
ure
(%)
1 6 7
Figure 1. Effect of watering treatments on soil moisture of pots containing single individuals of Agropyron repens and Lupinus perennis. Soil moisture was measured in all pots immediately before and one day after any watering event. Points represent the mean ± SE of eleven replicates over eight watering cycles (n=88). Orange bars indicate mean soil moisture field data at six and twelve days after a >1 cm rain event in experimental grasslands at Cedar Creek Ecosystem Science Reserve in east-central Minnesota (May-July, 2000-2010).
Agropyron
Lupinus
↓ Frequency↓ Volume
↓ Freq. / ↓ Vol.
Control
Control ↓ Freq. ↓ Vol. ↓ Freq./ ↓ Vol
0.0
0.5
1.0
1.5
2.0
2.5
3.0
3.5
Root
to S
hoot
Rati
o
Control ↓ Freq. ↓ Vol. ↓ Freq. / ↓ Vol.
0.0
0.5
1.0
1.5
2.0
2.5
3.0
3.5
Control ↓ Freq. ↓ Vol. ↓ Freq./ ↓ Vol
0.0
1.0
2.0
3.0
4.0
5.0
6.0
7.0
8.0
9.0
Tota
l Dry
Bio
mas
s (g) A
B
Control ↓ Freq. ↓ Vol. ↓ Freq./ ↓ Vol
0.0
1.0
2.0
3.0
4.0
5.0
6.0
7.0
8.0
9.0
Watering Treatment Watering Treatment
Figure 4. Biomass accumulation was significantly reduced in Lupinus but not in Agropyron under decreased water availability. The ratio of accumulated root to shoot biomass did not change in Lupinus, but there appeared to be a weak antagonistic effect between decreased volume and decreased frequency in Agropyron. Treatments not connected by the same letter are significantly different (p<0.05). Bars represent the mean ± SE of eleven replicates.
Agropyron Lupinus
Effects on Root, Shoot, and Total Biomass
AB A
B
B
C
A
A
B
AB AB A
Effects on Leaf Physiology and Water Use Efficiency
Watering Treatment
Watering Treatment
C
Figure 5. Percent changes in leaf physiology of plants grown with reduced water availability compared to the control. Effect on leaf net photosynthetic rates was variable (A); however, due to a decrease in conductance (B), the water use efficiency (CO2 assimilation per unit water loss) increased significantly in Agropyron under combined water stress conditions whereas Lupinus had a more variable response (C). Error bars represent standard error; asterisks indicate a significant difference between treatments and the control (** = p<0.05; * = p<0.10).
↓ Frequency ↓ Volume ↓ Freq. / ↓ Vol.
-90
-80
-70
-60
-50
-40
-30
-20
-10
0
10
% C
hang
e in
Con
duct
ance
↓ Frequency ↓ Volume ↓ Freq. / ↓ Vol.
-20
0
20
40
60
80
100
% C
hang
e in
Wat
er U
se E
ffici
ency
↓ Frequency ↓ Volume ↓ Freq. / ↓ Vol.
-70
-60
-50
-40
-30
-20
-10
0
10
20
30AgropyronLupinus
% C
hang
e in
Pho
tosy
nthe
sis
**
**
A B
CWatering Treatment
Watering Treatment
*
*
*
*
*
Water TreatmentsTable 1. Volume and frequency of water given to plants in each of four treatments.
Average days between watering
Volume per event (mL)
Control 5.7 300↓Frequency 11.4 600
↓Volume 5.7 150
↓Frequency / ↓Volume 11.4 300
Before watering
After watering
Measuring soil moisture
