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RESEARCH ARTICLE
Keywords Abstract
Climate change; ecosystem response;
lysimeter; mycorrhizae; rainfall pattern; soil Climate change scenarios for central Europe predict fewer but heavier rains
types. during the vegetation period without substantial changes in the total amount
of annual rainfall. To investigate the impact of rainfall patterns derived from
Correspondence
regionalised IPCC scenarios on agroecosystems in Austria, we conducted
J. Tabi Tataw, Institute of Zoology, University of
an experiment using 3 m2 lysimeters where prognosticated (progn.) rainfall
Natural Resources and Life Sciences Vienna,
A-1180 Vienna, Austria. patterns were compared with long-term current rainfall patterns on three
Email: h9740376@hotmail.com agriculturally important soil types (sandy calcaric phaeozem, gleyic phaeozem
and calcic chernozem). Lysimeters were cultivated with field peas (Pisum
Received: 2 July 2013; revised version sativum) according to good farming practice. Prognosticated rainfall patterns
accepted: 28 August 2013. decreased crop cover, net primary production (NPP) and crop yields, but
doi:10.1111/aab.12072 increased root production and tended to decrease mycorrhization. Soil types
affected the NPP, crop density and yields, weed biomass and composition, as
well as the root production with lowest values commonly found in sandy soils,
while other soil types showed almost similar effects. Significant interactions
between rainfall patterns and soil types were observed for the harvest index
(ratio crop yield versus straw), yield per crop plant, weed density and weed
community composition. Abundance of the insect pest pea moth (Cydia
nigricana) tended to be higher under progn. rainfall, but was unaffected by
soil types. These results show that (a) future rainfall patterns will substantially
affect various agroecosystem processes and crop production in the studied
region, and (b) the influence of different soil types in altering ecosystem
responses to climate change should be considered when attempting to scale-up
experimental results derived at the plot level to the landscape level.
reduces the mycorrhization rates (Augé, 2001; Porcel Table 1 Characteristics of the experimental soil types in the lysimeters
et al., 2003, Smith & Read, 2008) and increases insects (from soil analysis and partly from Steinitzer & Hoesch, 2005)
and invertebrate population (Ziska & Dukes, 2011).
Sandy Calcaric Gleyic Calcic
Weeds are expected to be less reduced by droughts Phaeozem Phaeozem Chernozem
than cultivated plants owing to their wide climatic or Parameters (S-soils ) (F-soils ) (T-soils )
environment tolerance, short generation time, small seed
Profile water content (mm) 250–500 400–700 460–730
size, uniparental reproduction capacity, high competitive
Infiltration (mm) 430 25 0
ability, high growth rate and phenotypic plasticity Evaporation (mm) 2800 3150 3150
(Whitney & Gabler, 2008; Clements & Ditommaso, 2011). pH value: CaCl2 7.4 7.6 7.6
The AMF has also been shown to alleviate drought Calcium carbonate (%) 0.143 0.260 0.106
(Augé, 2001; Porcel et al., 2003), improving the crop Phosphor, CAL (mg kg−1 ) 143 73 76
yield and water use efficiency (Bolandnazar et al., 2007), Potassium, CAL (mg kg−1 ) 187 246 286
Magnesium, available 83 273 277
while making plants less vulnerable to withstand various
(mg kg−1 )
abiotic stresses (Koltai & Kapulnik, 2010). As AMF Humus content 2.1 4.9 4.9
foster plants nutrient uptake it equally increases the Nitrogen, mineralisation 56 57 68
soluble protein content improving plants quality for (mg kg−1 7 days−1 )
herbivores (Subramanian & Charest, 1998). Reduction Boron, available (mg kg−1 ) 1.3 2.7 2.9
in precipitation increases insect mortality, affecting its Iron, EDTA (mg kg−1 ) 69 44 39
Manganese, EDTA 81 34 33
abundance, morphology and physiology (Moran et al.,
(mg kg−1 )
1987; Robinson et al., 2012). Copper, EDTA (mg kg−1 ) 3.3 3.4 3.2
Reduction in precipitation was tested on the soil Zinc, EDTA (mg kg−1 ) 4.6 4.6 4.7
types calcaric phaeozem (S-soils ), gleyic phaeozem (F-soils ) Sand (%) 67.9 21.5 22
and calcic chernozem (T-soils ), representing 80% of the Silt (%) 19 50 67 55
agricultural soil in Austria’s most fertile region (region Clay (%) 9.9 27.83 23
Cation exchange capacity, 11.29 25.13 26.00
of Marchfeld). S-soils are highly sandy, with very low
mmol/100
profile water and evaporation; F-soils have very high
clay content, highly mottled subsoils, with high profile CAL, calcium acetate-lactate method; EDTA, Ethylenediaminetetraacetic -
water and evaporation; while T-soils are highly silty acid.
with the highest profile water content (Table 1). Soil
types and its characteristics have been demonstrated to much lower groundcover and root biomasses, which are
affect several processes in agroecosystems, such as the indicators for soil water und nutrient availability. In this
availability and supply of water to plants (Passioura, study, we tested the effects of current long-term average
1991), respiration and soil temperature (Koizumi et al., rainfall patterns versus future prognosticated rainfall
1999), plant growth, vegetation cover and yield (Mako patterns based on regionalised global climate change
et al., 2008; Bestland et al., 2009; Genxu et al., 2009), the models simultaneously on three different soil types in
transfer and interaction of mineral nutrients (Echevarria a large-scale lysimeter facility.
et al., 2003; Matias et al., 2011), and the physical, chemical
and biochemical properties of soils (Rhoton et al., 1993;
Paz-Ferreiro et al., 2011), while higher soil sand content Materials and methods
has been shown to improve AMF colonisation (Zaller
Experimental site
et al., 2011).
Both the effect of precipitation and soil types on This experiment was carried out in 2011 using 18
agroecosystem processes have been studied in isolation; cylindrical steel (Cr/Ni 18/9) lysimeters each with
however, it is unclear how these important factors a surface area of 3.02 m2 and 2.45 m soil depth.
interact. On the basis of previous findings we hypothesize Lysimeters were located in Vienna, Austria and situated
that soil types with lower water holding capacities under a 10 m × 46 m tunnel covered with transparent
and/or nutrient availability like S-soils will interact polyethylene film (Fig. 1). Tunnels were open at the
with lower precipitation disrupting nutrient and water front and back and had 2-m-high openings at both length
transportation into and within the plants, slowing sides to allow proper ventilation.
down plant’s physiological activities like photosynthesis, The soil types (S-soils , F-soils and T-soils ) were filled each
subsequently reducing plants biomass, crop yields and into six lysimeters. These soil profiles were carefully
weeds abundance more than F-soils and T-soils with higher excavated from field sites and filled into the lysimeters
soil water content. Thus, S-soils are expected to have with their natural bulk density of 1.4 g cm−3 . Each soil
Measurements
65
60
55
50 Curr. rainfall
45 Progn. rainfall
Rainfall in mm
40
35
30
25
20
15
10
5
0
23 May
25 May
27 May
29 May
31 May
02 June
04 June
06 June
08 June
10 June
12 June
14 June
16 June
18 June
20 June
22 June
24 June
26 June
28 June
30 June
02 July
Date
Figure 2 Current and prognosticated rainfall amounts applied onto field pea stands during the vegetative period from May to July 2011.
within this area were successively removed, identified to test and Kolmogorov–Smirnov, respectively. Parameters
plant family level, counted and their mass weighed after that did not meet criteria for parametric tests were
drying at 50°C for 48 h; weeds growing on the remaining transformed using Boxcox transformations. Afterwards,
lysimeter area were pulled by hand and weighed. Total all parameters (total biomass, pea, straw, weed, harvest
weed biomass of each lysimeter was calculated by adding index, plant density, root production, mycorrhization,
the biomass of the permanent and the remaining plot root–shoot ratio, weed families and number of pea
area. moth) were analysed using a two factorial analysis of
Field pea plants and weeds were harvested by hand variance (ANOVA) with precipitation (two levels: curr.
cutting them 5 cm above the soil surface. Pea yield rainfall versus progn. rainfall) and soil types (three levels:
was obtained by threshing the sheets in the laboratory. F-soils , S-soils and T-soils ) as factors. We also performed
Field peas and straw were ground and N content was correlations between LAI and biomass (Spearman’s rank
determined using an elemental analyser (LECO TruMac, correlation coefficient) and pea yield and root production
St. Joseph, MI, USA). Crop P, K and Mg contents (Pearson’s correlation coefficient). All statistical analyses
were determined by inductively coupled plasma atomic were performed using the freely available software R
emission spectroscopy (ICP-AES, Thermo Scientific, iCAP (Free Software Foundation, Inc., Boston, MA, USA;
6000 series, Waltham, MA, USA). www.r-project.org).
No insecticide was used and insect pest population was
determined by direct sampling. At harvest abundance of
the Pea moth Cydia nigricana Fabricius (Lepidoptera: Tortri- Results
cidae) was counted in pea sheets on 10 randomly chosen Future rainfall pattern reduced the net primary produc-
crop individuals per lysimeter. Pea moth abundance per tion (NPP), weed abundance, pea biomass and yield more
m2 was calculated by multiplying the abundance plant−1 on S-soils than on T-soils and F-soils . The NPP, harvest index,
with crop density. pea biomass plant−1 , pea yield plant−1 , root produc-
tion and root-to-shoot ratio were significantly affected
by both rainfall and soil types (Table 3). The NPP under
Statistical analyses
progn. rainfall was 29% lower than under curr. rainfall
First, we tested the normal distribution and variance patterns. The NPP of S-soils was 36% lower compared
homogeneity of each parameter using the Shapiro with F-soils and 43% lower than T-soils , but insignificant
Table 2 Chemistry of the flow through water (leachate) on S-soils (except sample 25 on T-soil )
between F-soils and T-soils biomass (Fig. 3A). Groundcover Root growth was significantly different among the various
was significantly reduced under progn. rainfall, however soil types. Between all soil types the root-to-shoot ratio
unaffected by soil types; LAI was marginally significantly differentiated significantly (data not shown).
affected by rainfall and soil type (Table 3). The harvest index was 9% lower under progn. rainfall
Pea biomass, yield and weed biomass were significantly patterns than under current rainfall patterns. S-soils had
affected by soil types, but not by rainfall patterns (Table 3). the lowest harvest index and differentiated from the
Harvest index, pea yield per plant and weed density T-soils and F-soils by 39% and 42%, respectively, whereas
showed significant interactions between soil types and T-soils had 6% lower harvest index than F-soils (Fig. 4).
rainfall (Table 3). The pea and straw biomass production Root mycorrhizal colonisation rate was on average 22%;
in S-soils was 37% lower than F-soils and 35% lower than however, it was not affected by soil types; progn. rainfall
T-soils (Fig. 3B). The pea yield per m2 was significantly showed a trend towards lower mycorrhization rates
affected by soil type and marginally significantly affected compared with curr. rainfall (Table 3; Fig. 5).
by rainfall; S-soils produced the lowest pea yield being 63% Weed production was significantly affected by soil
lower compared with F-soils and 59% lower than T-soils ; types, with a trend towards decreasing weed production
F-soils and T- soils had similar yields (Table 3, Fig. 3C). under progn. rainfall (Table 3). S-soils had 50% less
Root production before implementing rainfall treatments weed biomass than T-soils and 34% less weed biomass
was significantly different between soil types (Table 3): than F-soils . Weed density was unaffected by rainfall
S-soils showed higher root production than F-soils and T-soils , or soil types (Fig. 6A). Weed communities consisted
whereas the root production between T-soils and F-soils was of the families Asteraceae, Chenopodiace, Polygonaceae and
similar (data not shown). One month after implementing Poaceae. The relative contribution of these families to
the rainfall treatments, root production was significantly the weed community was unaffected by rainfall or
affected by rainfall and soil types (Table 3; Fig. 3D). Across soil types (Fig. 6B), although there were considerable
all soil types the root growth under progn. rainfall was changes in the contribution of these families to the weed
on average 53% higher than under curr. rainfall patterns. communities.
Table 3 Analysis of variance results on effects of three different soil types (gleyic phaeozem – F-soils , sandy calcaric phaeozem – S-soils and calcic
chernozem – T-soils ) and rainfall patterns (curr. rainfall versus progn. rainfall) on agroecosystem variables in field peas
Variable F P F P F P
Ground cover (70 DAS, %) 1.359 0.294 16.474 0.002 0.556 0.588
LAI (90 DAS) 3.391 0.068 3.286 0.095 0.930 0.421
Net primary production (g m−2 ) 28.676 < 0.001 13.3 0.003 0.937 0.419
Pea + straw (g m−2 ) 4.492 0.035 2.951 0.111 0.754 0.491
Pea (g m−2 ) 12.486 0.001 3.979 0.069 0.081 0.922
Weed (g m−2 ) 9.602 0.003 2.692 0.127 0.228 0.800
Harvest index 119.093 < 0.001 5.837 0.033 12.285 0.001
Plant density (ind. m−2 ) 8.574 0.005
Biomass per plant (g) 31.522 < 0.001 11.093 0.006 1.616 0.239
Pea per plant (g) 71.842 < 0.001 17.567 0.001 4.459 0.036
Root production, pretreatment (g m−2 ) 16.590 < 0.001
Root production, treatment (g m−2 ) 11.769 0.001 8.438 0.013 0.449 0.648
Mycorrhization, pretreatment (%) 1.583 0.238
Mycorrhization treatment (%) 0.121 0.887 3.736 0.077 0.193 0.827
Root/shoot ratio 23.508 < 0.001 20.427 0.001 1.265 0.317
Weed density (ind. m−2 ) 0.863 0.447 0.000 0.987 3.775 0.053
Pea moth infestation (ind. m−2 ) 0.079 0.925 1.736 0.212 0.077 0.926
Across soil types, the abundance of pea moth (C. future rainfall patterns with a reduced ground cover and
nigricana) was on average 105% higher under progn. aboveground production but increased root production.
rainfall than under curr. rainfall; however, this was not The allocation of production into roots is probably a
statistically significant; soil types had no influence on C. stress reaction counteracting the induced drought by
nigricana (Fig. 7). increasing the root surface area of water absorption
Leaf area index significantly correlated with pea likewise extending deeper to meet the underground
biomass (r = 0.724, P = 0.024). There was no correlation available water (Masilionyte & Maiksteniene, 2011).
between mycorrhization rate and the pea yield, root We attribute the reduction of NPP under progn. rainfall
production or NPP (data not shown). patterns to differences in the soil profile water content,
Analysing the soil NH4 and NO3 contents from 0.1 M infiltration and evaporation rates (Steinitzer & Hoesch,
KCl soil extract showed strong increase in average 2005). In a lysimeter experiment with seven different
NO3 content from 0.416 to 2.225 μg g−1 on S-soil under crops including field pea, it was shown that the straw yield
prognosticated climate (Table 4). By the end of the responded positively to moisture with a 21% increase for
experiment no leachate was collected on F-soils and barely pea straw biomass under irrigation (Gan et al., 2009). The
one sample on T-soils , whereas on S-soils the leachate positive correlation between NPP and the LAI showed a
average NO3 content was almost twice as much with stronger effect of the climate on the vegetative growth and
progn. treatment, while NH4 and P contents were almost confirms findings that induced drought being responsible
the same for both treatments (Table 2). also for the reduction in crop cover rate (Cui & Nobel,
1992; Augé, 2001; Echevarria et al., 2003; Porcel et al.,
Discussion 2003; Matias et al., 2011). Decrease in harvest index
under progn. rainfall was in contradiction to the findings
Effects of rainfall patterns
of Martin & Jamieson (1996) associating increase in field
Simulated future rainfall patterns with 30% decreased pea harvest index to sensitivity in reproductive growth,
rainfall amount during the vegetation period and but is in conformity with the findings of others attributing
36% longer dry periods between rainfall events than it to photosynthetic changes (Sanchez et al., 2001).
the current long-term rainfall patterns affected several The observed increased root growth on all soil types
important processes within this agroecosystem. It was under progn. rainfall indicates that soil conditions in the
very interesting to observe most changes just about three soil types were still suitable for root extension
4 weeks after implementing treatments, which differ (Passioura, 1991; Feiziene et al., 2011). Overall root
by only 38 mm rainfall. Field pea stands responded to AMF colonisation was low, suggesting that AMF is not
A 700 B 400
600 350
Net Primary production (g/m2)
0 0
F S T F S T F S T F S T
Curr. rainfall Progn. rainfall Curr. rainfall Progn. rainfall
C 200 D 50
180 45
Root production (g d.wt./m2)
160 40
140 35
Pea yield (g/m2)
120 30
100 25
80 20
60 15
40 10
20 5
0 0
F S T F S T F S T F S T
Figure 3 Net primary production (A), biomass of field pea + straw (B), pea yield (C) and root production (D) in field peas at different soil types (gleyic
phaeozem – F-soils , sandy calcaric phaeozem – S-soils and calcic chernozem – T-soils ), under current and prognosticated rainfall patterns. Means ± SD,
n = 3.
very important in this leguminous crop. Nevertheless, a C. nigricana were collected under progn. rainfall than
trend towards reduced AMF colonisation under progn. under curr. rainfall. Although this difference was not
rainfall could be attributed to the fact that water statistically significant owing to high variation between
stress causes plants to be more metabolically perturbed. lysimeters, this indicates that pest species living in sheets
According to Augé (2001), the fungus strongly competes benefitted from future rainfall patterns. It has long been
for root allocates with the onset of stress, leading to known that pea moth is more abundant on pea varieties
reduced mycorrhization rates in response to resist drought with later flowering dates and longer flowering duration
stress (Stahl & Christensen, 1982; Cui & Nobel, 1992; (Nolte & Adam, 1962) and it could also be shown that
Subramanian & Charest, 1998; Augé, 2001; Bolandnazar the abundance of this pest species also correlates with the
et al., 2007). The reduced AMF trend observed on all soil pea cropping area in the surroundings (Thoeming et al.,
types with reduced progn. rainfall could be attributed 2011) as known for other crops (Zaller et al., 2008).
to reduced soil water content (Stahl & Christensen,
1982), contradicting the findings of Cui & Nobel (1992)
Effects of soil types
who associated higher colonisation with improved water
availability. The three soil types differed mainly in sand, silt, clay and
Overall, there were very few insect pests on the crops in humus contents, soil water capacity and cation exchange
the experimental year. Nevertheless, considerably more capacities. Overall, crops and weeds in sandy soils
1 50
45
0.8
40
35
0.6
Straw 30
Pea
0.4 25
20
0.2 15
10
0
F S T F S T 5
0
Curr. rainfall Progn. rainfall F S T F S T
A B
1400 Rest 100%
Poaceae
Proportion of weed families
1000 Chenopodiaceae
Asteraceae
800 60%
600 40%
400
20%
200
0 0%
F S T F S T F S T F S T
Figure 6 Absolute (A) and relative (B) abundance of weed families per m2 in field peas at different soil types (gleyic phaeozem – F-soils , sandy calcaric
phaeozem – S-soils and calcic chernozem – T-soils ), under current and prognosticated rainfall patterns. Means, n = 3.
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