The Effect of Rotation and In-Crop Weed Management on the Germinable Weed

Seedbank after 10 Years

Author(s) :Robert H. Gulden, Derek W. Lewis, Jane C. Froese, Rene C. Van Acker, Gary B. Martens,
Martin H. Entz, Doug A. Derksen, and Lindsay W. Bell
Source: Weed Science, 59(4):553-561. 2011.
Published By: Weed Science Society of America
DOI:
URL: http://www.bioone.org/doi/full/10.1614/WS-D-11-00001.1

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Weed Science 2011 59:553–561

The Effect of Rotation and In-Crop Weed Management on the Germinable Weed
Seedbank after 10 Years
Robert H. Gulden, Derek W. Lewis, Jane C. Froese, Rene C. Van Acker, Gary B. Martens, Martin H. Entz,
Doug A. Derksen, and Lindsay W. Bell*
Agricultural production systems that reduce the use of in-crop herbicides could greatly reduce risks of environmental
damage and the development of herbicide-resistant weeds. Few studies have investigated the long-term effects of in-crop
herbicide omissions on weed seedbank community size and structure. A crop-rotation study was sampled 10 yr after a
strictly annual rotation and an annual/perennial rotation were exposed to different in-crop herbicide omission treatments.
In-crop herbicides were applied either in all annual crops (control), omitted from oats only, or omitted from both flax and
oats. Seedbank densities were greatest when in-crop herbicides were omitted from flax and oats, and this treatment also
reduced crop yield. Shannon-Wiener diversity differed among crops in the annual crop rotation and among herbicide
omission treatments in the perennial rotation. Herbicide omissions changed the weed-community structure in flax and in
wheat and canola crops in the annual rotation enough to warrant alternate control methods in some treatments. The
magnitude of the effects on the seedbank parameters depended largely on the competitive ability of the crop in which
herbicides were omitted. No yield response to omitting herbicides in oats indicated that standard weed management
practices have reduced weed populations below yield-loss thresholds.
Nomenclature: Canola, Brassica napus L. BRSNN; oats, Avena sativa L. AVESA; flax, Linum usitatissimum L. LIUUS;
wheat, Triticum aestivum L. TRZAX.
Key words: Community assembly, crop rotation, herbicide omission, weed seedbank.

Agricultural production systems that reduce herbicide usage
are required to lower the risks of environmental damage and
to slow or prevent the onset of herbicide resistance in weed
populations. Herbicide resistance, especially after repeated use
of the same selective in-crop herbicides, is an increasing
problem in western Canada and elsewhere (Beckie et al. 2008;
Heap 1997). In response to these challenges, systems that
reward growers for using fewer chemicals, such as pesticide-
free production (PFP), have been developed (Schoofs et al.
2005). PFP avoids the use of in-crop herbicides and
preseeding or PRE soil residual herbicides, but allows
nonresidual fallow weed control, such as PRE and postharvest
application of herbicides like glyphosate. PFP encourages
the use of thresholds and, unlike the rigid rules of organic
production, allows farmers to decide to apply in-crop
herbicides from season to season tactically, depending on
weed pressure. Expected benefits from flexible production
systems such as PFP include a reduction in the selection
pressure for developing pesticide-resistant biotypes, reduced
input costs, a reduction in the pesticide load on the
environment, and reduced human (producers and consumers)
exposure to pesticides, without compromising productivity.
In 2000, a field experiment was initiated in the northern
Great Plains of Canada to investigate the impact of PFP in
oats or in both oats and flax in rotation with other annual
crops or with alfalfa (Medicago sativa L.) (Schoofs et al. 2005).
Over the first 4 yr of the experiment, total aboveground weed
population densities varied substantially. Reduced in-crop
DOI: 10.1614/WS-D-11-00001.1
* First, second, third, fifth, and sixth authors: Assistant Professor, Senior
Research Technician, Assistant Professor, Senior Lecturer, and Professor,
Department of Plant Science, 222 Agriculture Building, 66 Dafoe Road, University
of Manitoba, Winnipeg, MB R3T 2N2, Canada; fourth author: Professor,
Department of Plant Agriculture, Crop Science Building, University of Guelph, 50
Stone Road East, Guelph, ON N1G 2W1, Canada; seventh author: Research
Scientist, Brandon Research Centre, P.O. Box 1000A RR3, Agriculture and Agri-
Food Canada, Brandon MB R7A 5Y3, Canada; eighth author: Research Scientist,
CSIRO Sustainable Ecosystems/APSRU, P.O. Box 102, Toowoomba, Queensland
4350, Australia. Corresponding author’s E-mail: gulden@cc.umanitoba.ca
herbicide use increased weed densities in crops where
herbicides were not used; however, the effects of in-crop
herbicide omission on subsequent non-PFP crops were not
clear after 4 yr. Weed-community data were not reported in
Schoofs et al. (2005).
Weed management practices are strong filters that affect
weed-community size and structure. Midseason aboveground
weed communities are most influenced by in-crop weed
management practices (Barberi et al. 1997; Doucet et al.
1999) including the crop’s ability to interfere with weeds
(Leroux et al. 1996) and herbicide applications (Gulden et al.
2010). These are not independent, as in-crop herbicide
options are closely linked to crop species. Crop rotation also
can be used to reduce weed populations (reviewed in Liebman
and Dyck 1993) and alter aboveground weed-community
assembly (Légère et al. 2005; Swanton et al. 1999, 2006).
Time influences the relative contribution of factors that filter
the weed community. Swanton et al. (2006) reported greater
influence of management techniques on the weed-community
assembly in the early years of an experiment, but as the
rotation became more mature, crop species had a greater
influence. Weeds that pass through these management filters
by avoiding or tolerating management practices contribute
most to the seedbank.
Weed seedbanks have been described as a memory reflective
of the management history of a system (Cavers 1995), and
conform to assembly theory (Booth and Swanton 2002) by
being influenced by filters that affect community size and
composition. Weed seedbank densities are much greater and
are considered to be more stable than weed plant communities
(e.g., Sosnoskie et al. 2006), but have been studied less
frequently than weed plant communities because of labor and
space restrictions. In the northern Great Plains region, several
previous workers have examined the weed community in the
seedbank in cropping systems (Derksen and Watson 1998;
Harbuck et al. 2009; Mickelsen and Stougaard 2003; Thorne
et al. 2007). In organic systems where herbicides are not used,
seedbank densities tend to be greater (Albrecht 2005;

Gulden et al.: Weed seedbank response to reduced herbicide use N 553

Menalled et al. 2001), although this is not always the case
(Davis et al. 2005). Tillage influences seedbank densities
(Cardina et al. 2002; Feldman et al. 1997) with greater seed
numbers and a more diverse community composition as the
frequency of tillage declines. However, our knowledge of the
effects of herbicide omission on weed seedbank community
size and structure is limited.
The aim of this study was to investigate the weed seedbank
after 10 yr of reduced in-crop herbicide use in two crop
rotations, an annual crop rotation and an alfalfa-crop rotation.
In-crop herbicide use is an important management filter for
weed communities, and in-crop herbicide omissions were
expected to affect germinable weed seedbank parameters
such as seedbank density, seedbank diversity, and seedbank
community assembly. We hypothesized that weed-community
size increases and its structure changes in response to
decreased in-crop herbicide use. We also hypothesized that
in-crop herbicide omissions are more important filters for the
weed seedbank community than crop or rotation.
Materials and Methods
Field Experiment. In 2000, a crop-rotation experiment was
initiated to study the impacts of reduced in-crop herbicide use
at the Ian N. Morrison Research Farm located in Carman,
Manitoba, Canada (49u299480N, 98u 29260W, 267 m above
sea level). The experimental design, treatment structure, and
field operations were described in full by Schoofs et al. (2005).
In brief, the study was fully phased where all crops were
present each year, and consisted of regionally typical annual
crop rotation (flax–oats–canola–wheat) and an alfalfa-crop
rotation (flax–oats–alfalfa–alfalfa) arranged as a randomized
complete block design with three replicates. In each replicate,
each rotation was subjected to three treatments: the first where
in-crop herbicides were not applied during oat crops, the
second where in-crop herbicides were not applied during both
flax and oats crops, and the third a control rotation where
selective in-crop herbicides were applied during all annual
crops. Thus, there were 24 treatments per replicate and
individual experimental units were 4 by 12 m in size. Preplant
glyphosate was applied to all treatments at a dose of 1,332 g
active ingredient ha21 and the same dose was used to
terminate the alfalfa stand in late summer or early fall of the
second year, about 1 mo after the second cut of hay that
season. In-crop herbicides and application doses for each crop
are shown in Table 1. In the establishment year of alfalfa,
grassy weed control was applied only when deemed necessary,
which was in 2005, 2007, and 2008. The study was under no-
till management and the soil was disturbed only at the time of
planting. Crops were fertilized at the time of seeding based on
soil tests and recommendations. Each year, all crops were
planted on the same date (May 12, 2005, May 16, 2006, May
28, 2007, May 23, 2008). The planting dates fall into the
recommended time of planting for each crop for the region.
The preplant or PRE glyphosate applications occurred on
May 17, 2005, May 16, 2006, May 29, 2007, and May 21,
2008. Canola seed was treated with fungicide (trifloxystrobin,
metalaxyl) and insecticide (clothianidin, carbathiin). No
additional fungicides and insecticides were used or needed
between 2005 and 2008. In-crop herbicides were applied on
June 15, 2005, June 16 and 23, 2006, June 21, 2007, and
June 18 and 27, 2008. In 2006 and 2008, in-crop herbicides
554 N Weed Science 59, October–December 2011
Table 1. List of active ingredients and doses for each crop to which
postemergence herbicides were applied. Preplant glyphosate was applied to all
crops and second-year alfalfa was terminated with the use of glyphosate. All
herbicides were applied with carrier volume of 108 L ha21.
Crop Herbicide Formulation Dose
g ai ha21
Wheat Thifensulfuron–methyl 33.5% 10
Tribenuron–methyl 16.65% 5
Clodinafop–propargyl 240 g L21 56
Flax Sethoxydim 450 g L21 500
Bromoxynil 280 g L21 280
MCPA 280 g L21 280
Canola Glufosinate ammonium 150 g L21 500
Oats Thifensulfuron–methyl 33.5% 10
Tribenuron–methyl 16.65% 5
Alfalfa year 1 Sethoxydim 450 g L21 500
in flax were applied on a later date after flax reached the
appropriate growth stage.
Data Collected. In early May 2009, before any field
operations were conducted or weed seedling recruitment
had begun, 16 soil cores (6.4-cm diameter) were taken to a
depth of 15 cm in each plot. The core locations were chosen
randomly within each experimental unit, but were not
removed from within 1 m of the plot margins. All soil core
samples for each plot were bulked into one plastic tray and
trays were moved to the greenhouse for weed seed grow outs.
In the greenhouse, the soil was watered as necessary and
recruited seedlings were enumerated by species at regular
intervals. In cases where species were difficult to separate at
the seedling stage (e.g., Setaria, Brassica, and Sonchus), species
were grouped. After weed-seedling recruitment had ceased,
the soil was mixed occasionally to promote further weed
seedling recruitment until no further recruitment was
observed. The trays were stored at 220 C for 3 wk to break
dormancy between successive recruitment assays (Cardina and
Sparrow, 1996). Each of the recruitment assay took about 7 to
8 wk, and three assays were required before weed seedling
recruitment from the seedbank was exhausted. Remaining
seeds were not elutriated from the soil samples.
From 2005 to 2008, crop seed yield of annual crops in both
rotations was determined by harvesting each experimental unit
with a plot combine at maturity. The harvested samples were
air-dried, cleaned, and weighed, and seed yield was adjusted to
a constant moisture content for comparison of data among
years. Aboveground biomass was taken in late July or early
August in the second-year alfalfa crop, dried to constant
weight at 40 C, and weighed. Although first-year alfalfa was
cut to prevent weed seed additions to the seedbank, first-year
alfalfa yields were not recorded in 2005–2008.
Statistical Analysis. From the seedbank data, total germin-
able weed seed densities were determined by summing all
counts for each experimental unit and expressing these on an
area basis. For each experimental unit, the relative abundance
of each weed species was calculated as its proportional
contribution to the total germinable weed seed density. The
relative abundance values were used to determine Shannon-
Wiener diversity (Shannon and Weaver 1949). Species
richness was determined by summing the number of different
species that occurred in each plot.

Figure 1. Total germinable weed seedbank density in an annual-crop only (top)
and alfalfa-crop (bottom) rotation after 10 yr of in-crop herbicide omissions
during oats and flax and oats. Crop and herbicide-omission treatments followed
by different letters are statistically significantly different based on Fisher’s
Protected LSD means comparison conducted within each main effect. Bars
indicate standard errors of the means.
To determine the impact of in-crop herbicide omission and
crop species on total germinable weed seedbank density,
Shannon-Wiener diversity, and species richness, these data
were subjected to mixed-model analysis (PROC MIXED)
with the use of SAS (Littell et al. 1996). Prior to final analysis,
residuals were tested for normality and outliers were removed
with the use of Lund’s test (Lund 1975). In the analysis of
variance, fixed factors included herbicide omission and crop,
and repetition was considered random. Throughout this
manuscript, crop refers to the crop species that was grown in
2008. Data were analyzed within crop rotation (annual crop
rotation vs. alfalfa-crop rotation) because of an unbalanced
design with respect to crop and to minimize high-level
interactions. Treatment means were separated based on
Fisher’s Protected LSD (a 5 0.05) difference using the
pdmix 800 macro (Saxton 1998). To determine the effects of
rotation on weed seedbank density and diversity, a second
analysis was conducted using only data from crops common
to both rotations. A similar analysis was conducted for crop
yield, with the exception that year was included as a random
variable and replication was nested within year. A separate
analysis of variance was conducted for alfalfa yield with
herbicide omission repetition as the fixed variable and
repetition and year as random variables.
A univariate variance component analysis was conduced for
each weed species with the use of the Type 3 option in the
mixed procedure of SAS. The contribution of only main
Gulden et al.: Weed seedbank response to reduced herbicide use
factors (herbicide omission, crop, and rotation) and not
interactions among these were expressed as the proportion of
total Type 3 sums squares.
To determine treatment effects on qualitative character-
istics of the seedbank weed community, multivariate
analysis was conducted on relative abundance values within
crops that were present in both cropping systems (i.e., oats
and flax). A similar analysis was also conducted for all other
crops within each cropping system. To determine which
multivariate method was most appropriate, a detrended
canonical correspondence analysis was conducted initially,
and the length of gradient of the eigenvalues was examined
(Leps and Smilauer 1999). This analysis indicated that
redundancy analysis (RDA) was the most appropriate
method. To reduce potential bias of rare weed species,
data were log-transformed and rare weed species were
downweighted during RDA analysis using CANOCO 4.5
(Leps and Smilauer 1999). Herbicide-system and rotation-
treatment combinations were treated as independent
variables. Species densities were treated as dependent
variables and years, and replications were considered
covariables. Monte Carlo permutations were used to test
the significance of the first two canonical axes (Økland
2003). The treatment structure was nominal, and scaling
was focused on intersample distances. Biplots were
generated for each analysis with the use of CanoDraw
(Leps and Smilauer 1999). Four separate analyses were
conducted: one for all oat crops, one for all flax crops, one
for wheat and canola combined, and a multivariate analysis
separating weed communities in both years of alfalfa.
Results
Total Germinable Seedbank. After 10 yr of reduced in-crop
herbicide use, the total germinable seedbank density differed
among herbicide-use treatments and among crop species
(Figure 1). In the annual rotation, reducing the use of
in-crop herbicides increased the total germinable seedbank
population in all crops, with an approximate doubling in
seedbank densities when herbicides were omitted in both oats
and flax. In all herbicide-omission treatments in the annual
rotation, total germinable seedbank densities were greatest
after flax, with the total germinable seedbank densities similar
among other crops. The results were different in the alfalfa-
crop rotation (Figure 1, bottom); total germinable seedbank
densities were not affected by omitting in-crop herbicides in
oats only, but there was an increase when in-crop herbicides
were omitted in both flax and oats. When herbicides were not
used in both flax and oats crops of the alfalfa-crop rotations,
germinable seedbank densities were lower after second year
alfalfa, but were similar after flax, oats, and first-year alfalfa.
Seedbank Weed Species Diversity Indicators. Both the
previous crop and herbicide omission affected the number of
weed species in the seedbank in the annual crop rotation.
More species were present in the seedbank when in-crop
herbicides were omitted in both oats and flax, but there was
no change in the number of weed species when herbicide were
omitted during oats only (Figure 2). Among the crops, the
number of species in the seedbank was greatest after flax and
canola, and lowest after wheat. In the alfalfa-crop rotation, the
number of weed species present was unaffected by herbicide
N 555

Figure 2. Species richness in an annual crop only (top) and alfalfa-crop (bottom)
rotation after 10 yr of in-crop herbicide omissions during oats and flax and oats.
Crop and in-crop herbicide omission treatments followed by different letters
are statistically significantly different based on Fisher’s Protected LSD means
comparison conducted within each main effect. Bars indicate standard errors of
the means.
omission, but there was significantly greater species richness
after oats than after flax.
Seedbanks were dominated by Setaria species, yellow
woodsorrel (Oxalis stricta L.), and redroot pigweed (Amaran-
thus retroflexus L.), which accounted for an average of 86.2%
of all germinable seeds in the seedbank under all treatments
(Figure 3). The remainder of the seedbank was comprised
of eight other individual species and two species groups.
Variance component analysis indicated that of the three main
factors tested in this study (crop species, herbicide omission,
and rotation), the crop species grown in 2008 consistently
contributed most to the variation in total seed numbers
for each species among the treatments (i.e., more available
sums squared were partitioned to this factor). The relative
importance of herbicide omission and crop rotation on
seedbank density, however, was unique for each weed species
present. Crop rotation affected the seedbank densities of
redroot pigweed, yellow woodsorrel, shepherd’s-purse [Cap-
sella bursa-pastoris (L.) Medik.], and Sonchus species more
than in-crop herbicide omission. Conversely, for all other
species, herbicide omission affected seedbank densities more
than crop rotation. For the three major weed species, crop,
rotation, and herbicide omission accounted for 50% or more
of the total variation observed in density among the
treatments. For minor species, these main effects accounted
556 N Weed Science 59, October–December 2011
for only a very small portion of the variation (, 20%) in
density observed among the treatments.
In the annual rotation, weed species diversity was the same
under all herbicide treatments and differed only among crops.
Weed species diversity in the seedbank was greatest after
the canola crop (1.30 6 0.07) and lowest after the wheat
crop (0.73 6 0.08). Seedbank weed diversity after flax was
intermediate (1.06 6 0.08) and seedbank weed diversity after
oats (0.85 6 0.7) was similar to flax and wheat. In the alfalfa-
crop rotation, seedbank weed diversity was affected by
herbicide use only, with significantly lower diversity in the
treatment where in-crop herbicides were omitted in both flax
and oats (1.05 6 0.12) compared to the other two treatments
(1.30 6 0.7).
For the diversity and the multivariate analyses, it is
important to note that three species groups contained
multiple weed species because of difficulties in speciation
at the seedling stage. These were Setaria species [SETSP 5
green foxtail (S. viridis L.), yellow foxtail (S. glauca), and
barnyardgrass [Echinochloa crus-galli (L.) Beauv.]], Sonchus
species [SONSP 5 annual sowthistle (Sonchus oleraceous L.)
and perennial sowthistle (S. arvensis L.)], and Brassica species
[BRASP 5 volunteer canola (B. napus) and wild mustard
(Sinapis arvensis L.)].
Seedbank Community Assembly. Weed-community assem-
bly was analyzed based on relative abundance values to focus
on qualitative differences among weed communities while
precluding bias from quantitative differences. Seedbank weed
communities were analyzed over both rotations in flax and
oats; however, community assembly only differed significantly
after the flax crop (Figure 4, oat data not shown). Following
flax, the first canonical axis clearly separated weed commu-
nities based on crop rotation, and the second canonical axis
separated weed communities based on in-crop herbicide
omission. Within each rotation, weed seedbank communities
were most affected by whether flax was treated with herbicides
or not. In the annual crop rotation, higher relative abundance
of yellow woodsorrel in the seedbank was strongly correlated
with in-crop herbicide use on flax. Omitting in-crop herbicide
use in flax in the annual rotation was correlated with increased
seedbank relative abundance of Brassica species and Canada
thistle, and to a lesser extent wild buckwheat. In the alfalfa-crop
rotation following flax, higher relative abundances of Setaria
species and lambsquarters were correlated with omitting in-
crop herbicides during flax. Flax treated with in-crop herbicides
in this rotation was correlated with higher relative abundance of
redroot pigweed, and to a lesser degree with increases in
shepherd’s-purse, and annual and perennial sowthistle.
In crops that only occurred in one rotation, seedbanks were
analyzed for differences between crops and in-crop herbicide
omission. In the annual rotation, weed seedbank communities
were significantly different between wheat and canola
(Figure 5). The first canonical axis clearly separated weed
communities between these crops, and the second canonical
axis separated the weed communities based on in-crop
herbicide omission. In wheat, weed communities were
different under all three herbicide treatments. Higher relative
abundance of yellow woodsorrel was most strongly correlated
with wheat in the control rotation. Higher relative abundance
of Setaria species was correlated with wheat in rotations in
which in-crop herbicides were omitted in oats or oats and flax.
Figure 3. Mean relative abundance and proportion of total variance allocated to main effects (herbicide omission, crop, and rotation) for each weed species found in the
seedbank. Standard errors of the means are indicated.
In-crop herbicide omission affected seedbank weed-commu-
nity assembly after canola less than after wheat. In canola,
seedbank weed communities were most different when in-
crop herbicides were omitted in flax and oats. This correlated
with a higher relative abundance of lambsquarters and to a
lesser extent, perennial weeds. When in-crop herbicides were
applied in flax, weed communities were similar and had
higher relative abundance of Brassica species, redroot pigweed,
and smartweed. Weed seedbank community structure after
alfalfa was not affected significantly by year or in-crop
herbicide omission in oats and flax (data not shown).
In the multivariate analyses that showed different weed
seedbank community assembly in response to in-crop
herbicide omissions (i.e., flax in both rotations and canola
and wheat in the annual rotation), the first two canonical axes
explained a large proportion of the variation in the relative
abundance of weed species (78.5 and 81.7%) (Figures 5 and
6). These analyses also identified rough cinquefoil, wild oats,
and Sonchus species as more ubiquitous weed species whose
relative abundance was less influenced by crop, rotation, or
herbicide omission than other weeds.
Crop Yield. Crop yield was not affected by omitting in-crop
herbicides in oats in either crop rotation (Figure 6). An overall
Gulden et al.: Weed seedbank response to reduced herbicide use
yield decrease in annual crops was observed in both rotations
when in-crop herbicides were omitted in both flax and oats.
Among the annual crops, the cereal crops achieved the highest
yields and yield of flax was the lowest in both rotations.
The yield of canola was intermediate. An interaction between
crop yield and in-crop herbicide use was not observed in
either rotation (annual rotation P 5 0.3245, alfalfa rotation
P 5 0.068). Second-year alfalfa yield was not affected by in-
crop herbicide omissions and averaged 3591 kg ha21 (6 226)
from 2005 to 2008.
Discussion
Seedling recruitment measurements during the first 4 yr of
this study indicated increasing weed population densities with
reduced in-crop herbicide use (Schoofs et al. 2005), but high
variation was observed among years. This seedbank evaluation
after 10 yr of treatment implementation was very effective at
discerning quantitative and qualitative differences in the weed
seedbanks between crops, rotations, and in-crop herbicide
omission. The species observed in the seedbank in this
evaluation agree with those reported in the aboveground weed
community in Schoofs et al. (2005). The aboveground weed
community changed substantially in the first 4 yr of the study
N 557
Figure 4. Biplot of the association between higher relative abundance of each
weed species with crop rotation and herbicide omission treatments in flax crops.
Letter codes indicate rotation (Fa 5 annual, Fp 5 alfalfa crop) and number codes
indicate in-crop herbicide-omission treatments (0 5 control, 1 5 no in-crop
herbicides in oats, 2 5 no in-crop herbicides in oats and flax). Weed species codes
are indicated in Figure 3. The percentage of the data explained by each canonical
variable (i.e., axis of the biplot) is indicated.
(data not shown). In the first year, weed communities
immediately prior to in-crop herbicide application were
dominated by redroot pigweed (38%) and lambsquarters
(43%) but also included Setaria spp. (12%) and lambsquarters
(7%) (data not shown). A gradual shift over the next 4 yr, the
last time for which community information was available,
resulted in an aboveground weed-community structure similar
to seedbank observations in 2010. Based on relative densities,
the community was dominated by Setaria (40%) and redroot
pigweed (45%). Yellow woodsorrel was documented for
the first time (10%) with lambsquarters (7%), polygonum
species (6%), wild mustard (3%), and dandelion (Taraxacum
officinale G.H. Weber ex Wiggers) (3%) as minor species.
The overall impacts of reduced in-crop herbicide in
competitive crops such as oats appear to be of limited
statistical and biological effect on quantitative and qualitative
characteristics of the weed seedbank. Omission of in-crop
herbicides in oats only had a small effect on total germinable
seedbank densities, species diversity indicators, and the weed
seedbank community structure. Oats are a competitive, cool-
season cereal that interfere well with later-emerging Setaria
species (Fleck et al. 2009). The data from this study confirm
the logic that it is better to omit an in-crop herbicide during a
competitive crop rather than a less competitive one. It must be
noted that the crop effect in this study is comprised of short-
term (i.e., the fecundity of the weed community in the 2008
crop) and longer-term components (i.e., the effects of the
previous cropping sequence on weed seedling recruitment,
management, and weed fecundity) that cannot easily be
separated. The degree to which these affect our observations is
unique to each weed species and is influenced by the seedbank
longevity of the species.
558 N Weed Science 59, October–December 2011
Figure 5. Biplot of the association between higher relative abundance for each
weed species (with herbicide-omission treatments in wheat and canola crops in
the annual rotation. Letter codes indicate crop (Wh 5 wheat, Can 5 Canola) and
number codes indicate in-crop herbicide-use-intensity (0 5 control, 1 5 no in-
crop herbicides in oats, 2 5 no in-crop herbicides in oats and flax). Weed species
codes are indicated in Figure 3. The percentage of the data explained by each
canonical variable (i.e., axis of the biplot) is indicated.
Omission of in-crop herbicides in both flax and oats
significantly increased seedbank density and significantly
changed the structure of the seedbank community. This
effect was also evident in subsequent crops in the rotation.
A confounded treatment structure precluded the clear
isolation of herbicide omission in flax alone; however,
greater dissimilarity in weed seedbank attributes when
in-crop herbicides were omitted in both oats and flax
compared to oats alone suggests that the observed effects
were more likely related to the uncompetitive nature of flax
rather than further in-crop herbicide omission. Poor ability
of flax to compete with weeds is well documented (e.g.,
Lutman et al. 1996) and in uncompetitive crops herbicides
have been shown to be an important filter for weed density
(Derksen et al. 1995; Harker et al. 2005; Norsworthy 2008)
and community structure (e.g., Doucet et al. 1999). On the
other hand, growing a perennial crop such as alfalfa in
rotation significantly reduced the seedbank populations of
weeds under all herbicide-use intensities. Perennial forage
crops are known to reduce weed seedbanks composed
primarily of annual weed species effectively (e.g., Ominski
et al. 1999). The alfalfa crop was cut three times, once in
the first year and two times before stand termination in the
second year. Each cut was conducted before weeds were able
to return seeds to the seedbank, ensuring that even the
short-lived alfalfa stand of only 1.5 yr in this study was
effective at managing weed seedbanks. Including crops with
different life cycles and harvest regimes within a rotation to
manage weed populations is generally not exploited to its
fullest potential by producers.
Diversity indicators (species richness and Shannon-Wiener
index) did not differ substantially among treatments in this

Figure 6. Mean crop yield (2005–2008) of annual crops in an annual crop only
(top) and alfalfa-crop (bottom) rotation after 10 yr of in-crop herbicide omissions
during oats and flax and oats. Crop and in-crop herbicide omission treatments
followed by different letters are statistically significantly different based on
Fisher’s Protected LSD means comparison conducted within each main effect.
Bars indicate standard errors of the means.
study, and values of diversity indicators were similar to
aboveground weed-community diversity indicators reported
in other studies in agricultural systems (Cathcart et al. 2006;
Doucet et al. 1999; Légère et al. 2005). Diversity indices in
weed communities with relatively few species provide a
measure of community structure whereby lower Shannon-
Wiener values indicate greater dissimilarity in densities among
species (i.e., increased dominance of certain species) in
communities with similar species richness. This is a reflection
of the response of individual species to the selection pressures
exerted by the imposed management practices. The domi-
nance of Setaria species in many treatments and species
replacement (i.e., a change in presence/absence or relative
abundance of species among treatments with no change in
species richness) of major and minor weeds among the
treatments contributed to these observations. Harbuck et al.
(2009) showed that species with high relative abundance
tended to be indicator species for a particular management
system. We did not conduct an indicator species analysis for
the seedbank; however, our multivariate analysis indicated
that in some treatments, less abundant weeds were strongly
correlated with certain treatments (e.g., Brassica species in the
annual rotation in flax, in which in-crop herbicides were
omitted). This suggests that these less common weeds may
also be indicative of management history.
We saw evidence of dramatic shifts in relative abundance and
rank among species in response to treatments in this study that
Gulden et al.: Weed seedbank response to reduced herbicide use
could have important practical implications. For example, in the
alfalfa-crop rotation after flax, redroot pigweed replaced Setaria
species as the dominant weed species. This may adjust the
management strategies such as using herbicides with a different
mode of action in subsequent crops such as oats. Eleven
herbicides with a broad range of efficacies for managing redroot
pigweed in oats are registered in the region, but only one
herbicide option with poor efficacy is registered for the control of
Setaria species in oats (Anonymous 2010). Thus, in oats, redroot
pigweed as a dominant species is less problematic than Setaria
species. Changes in abundance of minor weed species is not likely
to affect management practices as long as these species remain a
minor component of the weed community.
Aboveground weed-community studies have shown a
positive relationship between weed density, species richness,
and diversity (Cathcart et al. 2006; Doucet et al. 1999;
Gulden et al. 2010). The same was not observed in the
seedbank in this study (data not shown). In fact, omission
of herbicides in oats and flax when rotated with alfalfa
substantially increased seedbank density, with a concomi-
tant decrease in species diversity resulting from increased
dominance of Setaria species (data not shown). Increased
weed diversity in the seedbank following the canola crop
may be related to the use of the nonselective herbicide
glufosinate in the glufosinate-resistant canola crop by
effectively controlling dominant weeds or the ability of
canola to compete with warm season weeds such as Setaria
and pigweed. Oats and alfalfa are competitive crops
(Ominski and Entz 2001; Willenborg et al. 2005) in which
herbicide choices (Anonymous 2010) and herbicide use
(Table 1) are limited. This contributed to the lack of
qualitative seedbank response to herbicide omission in spite
of substantial differences in seedbank densities among
herbicide-omission treatments and also indicates that oats
may be more resilient to rotation and management induced
changes in the structure of the weed seedbank community.
For alfalfa, the results are less clear as this crop followed
oats in the alfalfa-crop rotation and therefore the effect of
herbicide omission on weed-community structure may be
confounded by the preceding oat crop. These observations
and the variation in total germinable seedbank densities
among crops within each rotation support the notion that a
significant component of the seedbank in these systems is
transient. Conn et al. (2006) also reported a significant
transient component in agricultural seedbanks.
Although variance component analysis indicated that
individual weed species in the seedbank were influenced
primarily by crop and less by rotation, the clear separation of
the seedbank weed communities in flax and oats in the
different rotations indicates the importance of crop rotation in
defining weed seedbank community structure. In addition,
the crop effect on the seedbank in this study was confounded
by cropping sequence. The effect of crop rotation on weed
communities is well documented by others (Bellinder et al.
2003; Légère and Stevenson 2002; Ominski et al. 1999;
Unger et al. 1999). This rotation effect, in addition to a
significant transient portion of the seedbank (i.e., seeds with a
short residence life in the seedbank), illustrates that qualitative
aspects of the seedbank can change relatively quickly in
response to management filters. Despite this transient
component in agricultural seedbanks, their consistency over
time is also recognized. Sosnoskie et al. (2006) measured the
weed seedbank dynamics in a 35-yr-old tillage and rotation
N 559
experiment over a period of three consecutive years. Their
data showed high consistency in relative seed densities among
species over the 3 yr of study. Therefore, we feel confident
that our results are an accurate representation of the impacts
of the treatments after 10 yr.
There were only three wind-dispersed weed species in this
experiment (Canada thistle, perennial sowthistle, and dande-
lion) and these contributed on average less than 5% to the
total germinable weed seedbank population in those experi-
mental units in which they were present (data not shown).
The potential mobility of wind-dispersed seeds among
experimental units may confound the data; however, their
overall contribution to seedbank composition was sufficiently
low that this was not an important factor in this study.
In 2005, linola was replaced with a traditional flax cultivar
in both rotations. The inherent yield difference between linola
and flax precluded a combined yield analysis with the early
years of the study. Crop yields for the first 5 yr of this study
can be found in Schoofs et al. (2005) and tended to be similar
to those reported here. Interestingly, the increase in the total
seedbank density in the annual rotation in response to
omitting in-crop herbicides in oats did not affect crop yield in
the annual rotation. When analyzed within each crop (data
not shown), omission of in-crop herbicides in both flax and
oats also had no effect on canola and wheat yield in the annual
rotation and the yield of oats in the alfalfa rotation. These are
known as competitive crops with efficacious in-crop weed
management options (Anonymous 2010), and as a result,
productivity is more resilient to increased weed populations
and seedbank size. Taken together, this suggests that in these
extensive agricultural systems, the ‘‘perfect acre’’ philosophy
(Swanton et al. 2008) of complete weed control is flawed and
that current prophylactic weed management practices may
already be reducing weed populations below yield loss
thresholds in these systems. Higher seedbank densities
without a concomitant decrease in crop yield suggest greater
weed seedling recruitment after the application of selective,
nonresidual, in-crop herbicides, which would enable increased
weed fecundity with no effect on crop yield, as weed seedlings
recruited after the critical period.
Conclusion
Long-term rotation experiments that investigate the effects
of different in-crop herbicide omissions under different crop
rotations are rare. After 10 yr of treatment implementation,
the results here clearly showed that weed seedbanks conform
to assembly theory and are influenced by management filters.
The most recent crop in rotation was the most important filter
in this study. Rotation and in-crop herbicide omissions
influenced the weed seedbank in a species-specific manner,
which contributed to different weed-community assembly
between the two rotations. Weed-community dynamics are
driven by many intrinsic and extrinsic factors that do not act
independently. In this study, this was more obvious in minor
weed species, where the factors examined here explained little
of the variation in their seedbank density. A more thorough
understanding of these complex interactions is required for
weed-community assembly to progress from an observational
to a predictive science. The effects of crop, rotation, and in-
crop herbicide omissions on the weed seedbank community
size and structure were also of practical significance. Omitting
560 N Weed Science 59, October–December 2011
in-crop herbicides is better in competitive crops than less-
competitive crops, omitting in-crop herbicides in some crop
rotations does not necessarily increase overall weed density or
diversity, and in some cases, omitting in-crop herbicides may
shift species dominance to some that demand the use of
herbicides with a different mode of action or the use of other
or additional management strategies. The lack of a yield
response to higher weed-seedbank densities in the treatment
where in-crop herbicides were omitted from oats warrants
further investigation.
Acknowledgments
The authors wish to thank the University of Manitoba for funding
and the summer students for their technical assistance throughout this
study. We thank the Grains Research and Development Corporation,
Australia and the Australian Academy of Science for their support of
Lindsay Bell on his study tour to the University of Manitoba.
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Received January 3, 2011, and approved April 28, 2011.
N 561