Agriculture

Ecosystems &
Environment
ELSEVIER Agriculture, Ecosystems and Environment 56 (1995) 93-108

Crop-weed interference as influenced by a leguminous or synthetic

fertilizer nitrogen source: I. Doublecropping experiments with
crimson clover, sweet corn, and lambsquarters
Elizabeth Dyck *, Matt Liebman, M. Susan Erich
Department of Applied Ecology and Environmental Sciences, University of Maine, Orono, ME 04469-5722, USA
Accepted 7 September 1995

Abstract

Field experiments were undertaken to assess the effect of N source (incorporated legume residue vs. synthetic fertilizer) on
crop-weed interference. In a 2 year study, a doublecropping system was used in which a crimson clover ( Trifolium incarnatum
L. ) green manure was followed by a crop of sweet corn (Zea mays L.) grown alone or with lambsquarters (Chenopodium album
L.). Inclusion of several rates of ammonium nitrate fertilizer in the experiment allowed determination of the clover's N
equivalency value ( 55 kg N ha- ~) and contrast of the clover treatment with a comparable rate of N fertilizer addition (45 kg N
ha- ~). Soil NO3-N concentration in the experiment at one week after corn and lambsquarters planting was 52% lower in the
clover than the fertilizer treatment. Differences in nitrate levels between the two treatments tended to decrease at subsequent
sampling dates. At two weeks after emergence, drymatter accumulation of lambsquarters was 72% lower in the clover than the
fertilizer treatment and remained 39% lower at final harvest. In contrast, sweet corn biomass accumulation in the clover treatment
was 31% lower than in the fertilizer treatment at 2 weeks after emergence but recovered to levels attained in the fertilizer
treatment as the growing season progressed. As a result of reduced lambsquarters growth, loss of corn drymatter accumulation
to weed interference was 8% in the clover treatment as compared to 28% in the fertilizer treatment. Results of a second experiment
in which crimson clover was followed by lambsquarters grown alone also showed a weed suppressive effect of the legume N
source in comparison to use of fertilizer N. These experiments demonstrate that use of legume green manure has the potential to
reduce the need for herbicide as well as synthetic fertilizer applications in subsequent crops.

Keywords: Trifolium incarnatum L.; Zea mays L.; Chenopodium album L.; N fertilizer; Legume green manures

I. Introduction suggest that substitution of legume green manure for

Legume green manures have been assessed primarily
in terms of their ability to provide nitrogen (N) to a
subsequent crop (e.g. Hargrove, 1986; Hesterman,
1988). Relatively little research has focused on the
effect of a legume N source on crop-weed interference.
However, there are at least two areas of research that
* Corresponding author.
fertilizer N may reduce weed interference, and thus the
necessity for herbicide use, in the subsequent crop.
On one hand, chemical constituents of a legume
green manure or products of its decomposition could
inhibit weed growth. Aqueous and volatile extracts of
such legumes as sweetclover (Melilotus spp.), berseem
clover (Trifolium alexandrinum L.), crimson clover
(Trifolium incarnatum L.), and hairy vetch ( Vicia uil-

0167-8809/95 /$09.50 © 1995 Elsevier Science B.V. All rights reserved

SSDIO 1 6 7 - 8 8 0 9 ( 9 5 ) 0 0 6 4 3 - 5
94 E. Dyck et al. / Agriculture, Ecosystemsand Environment56 (1995) 93-108
losa Roth) have been shown to reduce germination and
early seedling growth in bioassay studies (McCalla and
Duley, 1948; White et al., 1989; Bradow and Connick,
1990). Moreover, sensitivity to allelochemicals
derived from legumes has been found to vary among
species; White et al. (1989), e.g. found water extracts
of crimson clover and hairy vetch to negatively affect
germination and radicle elongation of pitted morning
glory (Ipomoea lacunosa L.) and wild mustard (Bras-
sica kaber [D.C.] L.C. Wheeler) more than that of
corn (Zea mays L.). Such a differential effect, i.e.
greater suppression of weed than crop growth, is essen-
tial if the allelopathic potential of a legume green
manure were to be used successfully as a weed man-
agement strategy.
On the other hand, differences in temporal availa-
bility of N between synthetic and organic N sources
could lead to reduced interference from weeds in sys-
tems using a legume green manure. This theory (pre-
sented in detail by Liebman and Dyck, 1993) is based
on a synthesis of several research findings and assump-
tions. First, application of synthetic N fertilizer under
weed-infested conditions may be of greater benefit to
weed than crop growth and, as a result, may depress
crop yield (Okafor and De Datta, 1976; Carlson and
Hill, 1985; Liebman, 1989). This differential effect of
N fertilizer on crop and weed growth is probably linked
to the capacity of many arable weeds for early rapid
growth and N uptake (Haynes et al., 1991; Seibert and
Pearce, 1993), characteristics that confer a competitive
advantage in the use of the immediately available N
that occurs with synthetic fertilizer application. Sec-
ond, manipulation of the timing of N availability, i.e.
use of split applications of N fertilizer during the grow-
ing season rather than as a single application at planting,
has been shown to benefit crop growth under weed-
infested conditions (Alkaemper et al., 1979). And
third, legume residue, in comparison to synthetic N
fertilizer, has been characterized as a form of slow or
delayed release N (Ladd et al., 1983; Mueller, 1987).
In this context, use of a legume N source may decrease
weed interference with crop growth through a slow
release of N, which could deprive weeds of an early
competitive advantage while benefiting those crops
whose maximal rates of growth and N uptake occur
later in the growing season.
Previous work on the effect of legumes on weed
management has largely focused on legumes as
smother crops (Nelson et al., 1991; Samson, 1991 ) or
as mulches in no-tillage systems (Janke et al., 1989;
Hoffman et al., 1993). In these studies, legume treat-
ments have been compared to non-leguminous cover
crops or to control treatments unamended with cover
crop residue. However, to determine the effect of a
legume as a N source on crop-weed interference, it is
more appropriate to compare use of the legume green
manure to that of synthetic fertilizer N.
Accordingly, a series of experiments was conducted
to determine the effect of N source (legume green
manure vs. N fertilizer) on weed and crop growth.
Because the investigators' interest was on the effect of
hypothesized qualitative differences between N
sources and because amount of N can also influence
crop-weed interactions (Lawson and Wiseman, 1977;
Carlson and Hill, 1985), a legume green manure treat-
ment was compared with an application rate of N fer-
tilizer that supplied an equivalent amount of N to the
subsequent test crop. It was hypothesized that the leg-
ume N source would result in ( 1 ) reduced weed growth
and (2) enhanced crop growth and yield under weedy
conditions in comparison to application of fertilizer N.
2. Methods and materials
2.1. Experimental system
The above hypotheses were tested within a double-
cropping system of crimson clover followed by sweet
corn. The doublecropping system was used to facilitate
temporal replication of the experiment. Crimson clo-
ver, which is capable of rapid biomass accumulation
(Knight and Hoveland, 1985), was selected as the
green manure crop to optimize biomass production
within the doublecropping system. Corn was used as
the test crop because corn production systems are heav-
ily dependent both on N fertilizer and herbicides (Eco-
nomic Research Service, 1993). Sweet corn was used
in particular because it had performed well in past sea-
sons at the experiment station following early summer
plowdown of overwintering legume green manures,
e.g. alfalfa (Medicago sativa L.) and red clover (Tri-
folium pratense L.). To further accommodate the con-
straints of a double-cropping system, a short season
variety (70 days) of sweet corn was used. Lambsquart-
ers (Chenopodium album L.), a major pest of corn
 E. Dyck et aL /Agriculture, Ecosystems and Environment 56 (1995) 93-108
Table 1
Dates of major field operations in Experiments 1 and 2
Field operation Experiment 1
1989
Crimson clover planted 11 May
Crimson clover incorporated 10 July
Sweet corn and lambsquarters planted 12-13 July
Experiment irrigated 22 July
First weeding 4--10 August
Second weeding 29-31 August
~'Lambsquarters only was planted in Experiment 2.
(Holm et al., 1977), was chosen as the infesting weed
species.
Although most recent studies of the effect of legume
residues on weeds have been conducted within no-
tillage systems, in these experiments the effect of incor-
porated legume residue was studied for two reasons.
First soil incorporation of residue has been shown to
accelerate or intensify microbially mediated transfor-
mations of N (e.g. immobilization (Varco et al., 1993)
and rate of N release from plant tissue (Wilson and
Hargrove, 1986)) that can in turn affect subsequent
plant growth. Second, incorporation, particularly shal-
low incorporation, serves to concentrate residue in the
seed and seedling rooting zones, which may intensify
effects of residue on early plant growth. To further
ensure crop and weed interaction with the legume green
manure, in these experiments crop and weed seeds were
planted into freshly incorporated legume residue.
Two experiments were undertaken. In Experiment 1
(conducted for 2 years), crimson clover was followed
by sweet corn grown alone and in association with
lambsquarters. To allow separation of direct effects of
N source on weed growth from that of crop interfer-
ence, Experiment 2 (conducted for 1 year) was under-
taken in which crimson clover was followed by
lambsquarters grown alone.
2.2. Site description, experimental design, and field
operations
Experiment 1 was initiated in 1989 at the University
of Maine Sustainable Agriculture Research Center in
Stillwater, Maine. The site consisted of both a Adams
fine sandy loam (sandy, mixed, frigid Typic Haplor-
95
Experiment 2
1990 1989
2 May 22 May
10 July 24 July
11 July 26 July a
16 July I August
2-4 August 18-23 August
15-16 August 14-15 September
thods) and a Boothbay silt loam (fine, silty, mixed,
frigid aquic Dystric Eutrochrepts), and the experiment
was blocked accordingly. In the 5 years before the start
of experimentation, a series of field crops of small
grains and legumes had been grown on the site. In 1990
the experiment was repeated on the same site used in
1989, each plot receiving the same experimental treat-
ment in both years.
The experiment used a split-plot randomized block
design with four replications. Main plots consisted of
six N treatments: crimson clover residue, a 0 N treat-
ment (receiving neither residue nor N fertilizer), and
ammonium nitrate applied at rates of 45, 90, 135, and
180 kg N ha- 1. The 0 N treatment and various fertilizer
rates were included to allow estimation of a N fertilizer
equivalency value of the crimson clover residue. Sub-
plots were weed-free corn or corn grown with lambs-
quarters. Plots measured 3.0m × 9.1 m to accommodate
four corn rows spaced 76 cm apart.
Specific dates of major field operations are given in
Table 1. In early May of each year the entire site was
rotovated once. Crimson clover (cv. Dixie) seed inoc-
ulated with Rhizobium trifolii was then sown in appro-
priate plots by hand at a rate of 84 kg ha J (a high
seeding rate was used to ensure sufficient clover stand
development within the doublecropping system). Non-
legume treatments were kept in bare fallow by roto-
vating twice. At onset of flowering, the clover was
mowed and its residue incorporated with a rotovator to
a depth of 10-15 cm. After treating all plots with P and
K fertilizer at rates recommended by the University of
Maine Soil Testing Lab (in 1989, 84 kg ha -1 each of
P205 and K20; in 1990, 84 kg ha -~ of P205 and 196
kg ha- 1 of K20), the entire experiment was disk har-
96 E. Dyck et al./ Agriculture, Ecosystems and Environment 56 (1995) 93-108
rowed (in 1989) or rotovated (in 1990). Within 2 days
of clover incorporation, sweet corn (cv. Sugar Buns)
was sown in all plots. Plots were heavily seeded with
corn in both years and then thinned to assure adequate
stands. In 1989, corn was planted mechanically. In
1990, in an attempt to achieve more even spacing of
corn plants within the row, corn was seeded by hand.
In each year, on the same day as corn planting ammo-
nium nitrate treatments were broadcast into appropriate
plots and then shallowly raked into the soil to a depth
of 2-3 cm. Lambsquarters seed (collected from the
experimental farm the previous year and acid-pre-
treated to remove hardseededness (Ellis et al., 1985) )
was then sown by hand over the corn rows in appro-
priate plots at a rate of 1.1 g m - 2. In 1989, clover plots
were seeded with double the amount of lambsquarters
seed used in the 0 N and N fertilizer treatments in
anticipation of possible inhibitory effects of the residue
on emergence. In 1990, equal amounts of lambsquart-
ers seed were sown in all weedy corn treatments in an
effort to document possible residue effects on emer-
gence. To ensure adequate crop and weed establish-
ment, plots were irrigated in both years with overhead
sprinklers within 12 days after planting; total irrigation
water delivered was 28 mm in 1989 and 21 mm in
1990.
Plots were handweeded and hoed twice in each sea-
son to remove all weeds from the corn alone treatment
and all weeds but lambsquarters from the corn +
lambsquarters treatment. To simulate field conditions
in which mechanical cultivation is used for weed con-
trol, iambsquarters plants in the weedy plots were con-
fined to bands 30 cm wide centered on the corn rows.
Corn and lambsquarters were thinned between 1-2
weeks after corn emergence. Target densities were 6.5
plants m -2 and 26 plants m -2 for corn and lambs-
quarters, respectively. The corn density is in accor-
dance with the standard planting practices for sweet
corn in the northeastern U.S. (Erhardt, 1991). The
weed density required to cause substantial crop yield
loss can vary between sites and years; to insure a com-
petitive interaction between crop and weed, the target
lambsquarters density selected was considerably higher
that the 8.7 lambsquarters plants m -2 reported by
Beckett et al. (1988) to produce significant yield loss
in field corn.
Experiment 2 was conducted in 1989 on a Boothbay
silt loam at a site adjacent to Experiment 1. A corn-
pletely randomized design was used with four replica-
tions. Nitrogen treatments were identical to those used
in Experiment 1, while plot size was 3.0 m X 4.6 m.
Dates of major field operations are given in Table 1.
Field operations and plot management techniques were
the same as those used in Experiment 1, with the fol-
lowing exception. Although lambsquarters stands were
kept free from other weeds throughout the growing
season, no attempt was made to thin the lambsquarters
stands in each plot to a constant density. Instead, counts
of plant number were made at each sampling date to
check for potential differences in density between plots
or treatments.
2.3. Sampling and analytical procedures
In both Experiments 1 and 2, crimson clover biomass
was sampled immediately before clover incorporation.
Shoot material was collected in each legume plot from
two 0.05 m 2 quadrats. In Experiment 1, emergence
counts of sweet corn, lambsquarters, and other weeds
were taken each year before thinning and weeding
occurred using five 0.23 m 2 quadrats per plot. Above-
ground corn and lambsquarters biomass were sampled
at approximately 2, 4, and 6 weeks after corn emer-
gence from the two center corn rows of each plot using
two 0.33 m 2 quadrats. At the end of the growing season,
six 0.33 m 2 quadrats were collected. Four of the quad-
rats were bulked and the freshweight of each species
determined. The remaining two quadrats were bulked
and both freshweight and dryweights determined in
order to generate a conversion factor of percent dry-
matter for each species on a plot by plot basis. At each
biomass harvest, heights of sampled corn and lambs-
quarters plants were determined to the furthest
extended leaf tip. Stem diameter measurements (taken
slightly above the soil surface) were made of corn
plants at all biomass harvests and of lambsquarters
plants at the final three harvests. Sweet corn and lambs-
quarters leaf number were determined at the first har-
vest date. In Experiment 2, lambsquarters biomass was
sampled at five dates during the growing season using
two 0.167 m 2 quadrats. Height and stem diameter meas-
urements and leaf counts were taken as in Experiment
1.
In both experiments, samples used in dryweight
determination were dried for several days at approxi-
mately 65°C before weighing. To determine whole
 E. Dyck et al. / Agriculture, Ecosystemsand Environment56 (1995) 93-108
Table 2
Crimson cloverbiomassaccumulation, N concentration,and N fertilizerequivalencyvalues in Experiment 1
Year Aboveground N concentration
drymatteraccumulation (kg ha- ~) (g kg - 1)
1989 5136 24.0
1990 4758 24.6
~Wherey is the total N accumulationin weedfreecorn and x is the N fertilizerrate.
plant N content, dried plant tissue samples were ground
to pass through a 40-mesh screen and subjected to the
block digestion procedure outlined by Wall and Gehrke
(1975). The digests were analyzed for NH4+ on a
Lachat automated ion analyzer.
Soil samples were collected at four dates after corn
planting in both years of Experiment 1. Using randomly
generated coordinates, five soil cores, taken to a depth
of 25 cm, were collected and bulked to form a com-
posite sample for each experimental plot. The samples
were immediately frozen and stored until they could be
extracted. Samples were extracted using 5 g (oven dry-
weight basis) soil/50 ml 1N KCL and a shake time of
30 min. Soil extracts were analyzed for both NH4-N
and NO3-N content on a Lachat automated ion analyzer.
2.4. Statistical analysis
Total crop N accumulation (drymatter accumula-
tion × crop N concentration), which is a more sensitive
indicator of external N supply than yield (Hargrove,
1986; Hesterman et al., 1992), was used as a basis for
estimating an N fertilizer equivalency value for crimson
clover. In each year of Experiment 1, a fertilizer equiv-
alency value was calculated using the following steps:
( 1 ) total N accumulation of corn grown under weed-
free conditions in the non-legume treatments was ana-
lyzed using polynomial contrasts for significant linear,
quadratic, or cubic trends, (2) the appropriate model
was then used to generate a prediction equation by
regressing mean treatment values of corn N accumu-
lation on N fertilizer rate, and (3) the mean value of N
accumulation in weedfree corn following crimson clo-
ver was inserted into the equation and an estimate of N
fertilizer equivalency of crimson clover calculated. The
crimson clover treatment was then compared to the N
fertilizer treatment closest to its equivalency value
using single degree of freedom contrasts within an anal-
ysis of variance. Data generated in Experiments 1 and
97
Predictionequationa r2 CalculatedN fertilizer
equivalencyvalue (kg N ha- ~)
y = 0.009x +5.507 0.58 52
y = 0.026x +9.070 0.83 58
2 were analyzed using the repeated measures option in
the GLM procedure of SAS (1990). Results are
reported as between effects (treatments averaged
across sampling dates) and within effects (treatment
by sampling date interactions). Multivariate analysis
was used to test hypotheses concerning sampling date
by treatment interactions.
3. Results and discussion
3.1. Experiment1
3.1.1. Crimson clover biornass accumulation and N
fertilizer equivalency values
Crimson clover aboveground biomass accumulation
averaged 4947 kg ha- 1 over both years of Experiment
1 (Table 2). The average N concentration of crimson
clover shoots was 24.3 g kg - t. In both years of the
experiment, trend analysis data indicated a significant
linear relationship between N accumulation in weed-
free corn and N fertilizer rate ( P < 0 . 0 5 ) . Prediction
equations generated from subsequent regression anal-
ysis provided estimates of N fertilizer equivalency as
ammonium nitrate for crimson clover of 52 and 58 kg
N ha - t , respectively, in 1989 and 1990 (Table2).
Nitrogen fertilizer equivalency values of legumes for
sweet corn have apparently not been previously cal-
culated. However, the values for sweet corn found in
this study appear reasonable in light of average values
for N contribution of crimson clover reported in studies
for no-tillage field corn systems in the southeastern
USA (72 kg N ha -1 (Smith et al., 1987)) and in
Maryland (63 kg N h a - t ; Holderbaum et al., 1990).
In subsequent analysis of soil, lambsquarters, and sweet
corn data, the crimson clover treatment was therefore
contrasted with the 45 kg N h a - t treatment, which in
both 1989 and 1990 was the N rate closest to the clov-
er's estimated N fertilizer equivalency value. For pur-
98 E. Dyck et al./ Agriculture, Ecosystems and Environment 56 (1995) 93-108
poses of comparison, results of the 0 N treatment have
also been reported.
3.1.2. Soil inorganic N
Because of divergent sampling dates between years,
soil data for each year of Experiment 1 were analyzed
separately. Temporal patterns of plant-available N in
the soil differed significantly between N sources (45
kg N h a - ~ and crimson clover) in both 1989 and 1990
(Table 3 and Table 4). In 1989, initial NO3-N and
NH4-N concentrations were 65% lower in the clover
than the fertilizer treatment. At subsequent sampling
dates, differences in nitrate levels between the two
treatments steadily decreased while ammonium con-
centration in the two treatments differed by less than 1
mg kg - 1. In 1990, NO3-N concentration in the legume
treatment was 40% lower than in the fertilizer treatment
at eight days after planting (DAP). By 59 DAP, how-
ever, nitrate concentrations in the clover treatment were
twice those in the fertilizer treatment. Levels of NH4-
N in 1990 did not differ between the two treatments.
Concentration of NHa-N was not affected by the
presence of weeds in either year (Table 4). However,
in both years, NO3-N concentrations were reduced in
all weedy treatments as the growing season progressed:
weed presence had lowered nitrate levels 42% at 53
DAP in 1989 and 67% at 59 DAP in 1990. No signifi-
cant differences in the effect of weeds on NO3-N levels
were detected between clover and fertilizer treatments
(N source X weed effects) in either year. In both years,
differences in total soil mineral N (NO3-N + NH4-N)
between clover and fertilizer treatments paralleled dif-
ferences in NO3-N concentrations in terms of both
weed and N source effects.
3.1.3. Emergence of lambsquarters, other weeds, and
sweet corn
In both years of Experiment 1, weed emergence in
the clover treatment was delayed or suppressed when
compared to emergence in the N fertilizer treatment.
Although crimson clover plots had been sown with a
higher rate of lambsquarters seed in 1989, lambsquart-
ers emergence did not differ between clover and fertil-
izer treatments (Table 5). In 1990, when an equal rate
of lambsquarters seed was sown in all treatments,
lambsquarters emergence in the clover treatment was
34% lower than in the fertilizer treatment at 13 DAP,
although by 21 DAP lambsquarters densities were
equivalent in the two treatments (Table 5 ). Emergence
of weeds other than lambsquarters (of which the maj or
species were Amaranthus retroflexus L. and Capsella
bursa-pastoris [L.] Medic.) was strongly suppressed
in the crimson clover treatment in both years: counts
of other weeds were 60% lower in the clover than the
fertilizer treatment at 18 DAP in 1989 and 74% lower
at the single count of other weeds taken at 13 DAP in
1990. Emergence data also indicate a possible inhibi-
tory effect of crimson clover on sweet corn emergence.
Although corn emergence in 1989 did not significantly
differ between N source treatments, in 1990 sweet corn
emergence in the clover treatment was 29% lower than
in the N fertilizer treatment at 13 DAP.
3.1.4. Lambsquarters and sweet corn stand densities
Lambsquarters densities at final harvest were 35
plants m - 2 in 1989 and 81 plants m - 2 in 1990. Lambs-
quarters number exceeded the target density in both
years because of flushes of emergence that occurred
after thinning. Despite increase in density after stand
establishment, analysis of lambsquarters counts taken
at each biomass harvest showed no difference in density
between the crimson clover and fertilizer treatments
(data not shown). After thinning, sweet corn density
was 6.8 plants m - 2 in 1989 and 7.3 plants m - 2 in 1990.
3.1.5. Lambsquarters and sweet corn growth
3.1.5.1. Seasonal effects on lambsquarters and corn
growth
Despite the use of a short season cultivar, sweet corn
failed to develop marketable ears before killing frosts
in either 1989 and 1990. Treatment effects on lambs-
quarters and corn growth have been analyzed in terms
of aboveground biomass accumulation,, height and
stem diameter development, and leaf number. Similar
sampling dates and comparable N fertilizer equiva-
lency values in each year allowed 1989 and 1990 plant
growth data to be analyzed jointly.
Although a joint analysis of 1989 and 1990 lambs-
quarters and sweet corn biomass accumulation data
showed a significant year effect, there was no evidence
of an interaction of year with N source or related effects
(year X N source × weed, year × N source × sampling
date, year X N source X weed X sampling date). Anal-
ysis of lambsquarters and sweet corn stem diameter and
height data produced similar results. Because of this
 E. Dyck et aL /Agriculture, Ecosystems and Environment 56 (1995) 93-108 99

Table 3
Soil inorganic N concentrations (mg N k g - ~ soil) at four sampling dates in 1989 and 1990 in Experiment 1

1989

N treatment 9 DAP a 29 DAP 53 DAP 84 DAP

Weedfree corn Weedy corn Weedfree corn Weedy corn Weedfree corn Weedy corn Weedfree corn Weedy corn

NO3-N
0N 15.4 14.6 29.4 19.1 9.6 4.1 2.9 1.2
45 kg N h a - ~ 26.8 25.8 28.6 34.5 16.2 8.4 5.4 1.8
Crimson clover 8.6 9.5 12.7 17.0 8.3 5.9 2.8 2.7
NH4-N
0N 1.7 1.1 1.3 1.4 0.8 0.6 2.7 2.7
45 kg N h a - l 8.9 7.0 0.7 0.8 0.8 0.7 2.7 2.7
Crimson clover 2.3 3.2 1.3 1.4 0.8 0.8 2.4 3.3

1990

N treatment 8 DAP 22 DAP b 37 DAP 59 DAP

Weedfree corn Weedy corn Weedfree corn Weedy corn Weedfree corn Weedy corn Weedfree corn Weedy corn

NO3-N
0N 9.3 9.7 11.6 10.8 6.6 6.1 1.3 0.5
45 kg N ha - l 16.9 20.0 21.1 21.0 11.2 8.9 2.7 1.1
Crimson clover 10,8 11,5 17.1 18.1 11.5 11.2 6.6 1.5
NH4-N
0N 1,4 1.2 34.6 33.6 1.1 1.1 1.1 1.2
45 kg N h a - ~ 7.3 5.4 36.0 20.7 1.8 0.8 1.2 1.1
Crimson clover 4.1 3.0 35.7 30.1 1.0 1.0 1.6 1.4

aDays after planting.~The high levels of NI-~-N occurring at 22 DAP in 1990 are a likely result of a heavy rainfall and subsequent flooding of
the experimental site on 16 DAP, which may have precipitated a flush of ammonification and temporarily impeded nitrifying activity.

similarity in N source effect between years, results from
a single year (1989) have been used to illustrate N
source x sampling date effects (Fig. 1).
3.1.5.2. Lambsquartersgrowth
Use of the crimson clover treatment substantially
reduced lambsquarters growth in the weedy corn plots
when compared to that of N fertilizer (Table 6). The
suppressive effect of the clover treatment was imme-
diate and strong: at the first sampling date (approxi-
mately two weeks after corn and lambsquarters
emergence) in 1989 and 1990, lambsquarters biomass
in the crimson clover treatment was 64% and 81%
lower, respectively, than in the fertilizer treatment.
Lambsquarters growth was also characterized by a sig-
nificant N source X sampling date interaction
(Table 6): in both years drymatter accumulation in the
clover treatment gradually increased in relation to that
in the fertilizer treatment after the first sampling date,
as can be seen when biomass is plotted on a natural log
scale (Fig. la). However, despite decreased differ-
ences between treatments as the growing season pro-
gressed, lambsquarters remained 37% and 42% lower,
respectively, in the clover than the fertilizer treatment
at final sampling dates in 1989 and 1990. In both years
lambsquarters growth in the clover treatment never
exceeded that in the 0 N treatment (Table 6).
Lambsquarters plants grown with crimson clover
consistently had both smaller stem diameters
(P<0.0001) and were shorter (P<0.0001) than
plants grown with N fertilizer (Fig. lb and lc). Both
lambsquarters stem diameter and height showed an
interaction of N source with sampling date (P < 0.05).
However, while the difference in stem diameter
between N source treatments steadily decreased with
100 E. Dyck et al. / Agriculture, Ecosystems and Environment 56 (1995) 93-108

Table 4
Results of multivariate analysis of variance for repeated measures on soil inorganic N levels at four sampling dates in 1989 and 1990 in
Experiment 1"

1989 1990

NH4-N NO3-N NH,,-N + NO3-N NH4-N NOa-N NH4-N + NOa-N

Between effects
N source (45 kg N - ~vs. crimson clover) ns * ** ns ns ns
Weed b ns **** **** ns **** ****
N source × weed ns ns ns ns ns ns
Within effects
N source × sampling date *** * ** ns ** **
Weed × sampling date ns *** *** ns ** * * ** * *
N source × weed × sampling date ns ns ns ns ns ns

~Before analysis, data were In-transformed to reduce heterogeneity of variances among sampling dates.bWeed signifies presence or absence of
lambsquarters.Significance levels: ****, P < 0.0001; ***, P < 0.001; * *, P < 0.01; *, P < 0.05; ns, not significant.

Table 5
Emergence of sweet corn, lambsquarters, and other weeds (plants m -2) at two sampling dates in 1989 and 1990 in Experiment 1

N treatment 1989 1990

12 DAPa 18 DAP ! 3 DAP 21 DAPb

Sweet Lambs- Other Sweet Lambs- Other Sweet Lambs- Other Sweet Lambs- Other
corn quarters weeds corn quarters weeds corn quarters weeds corn quarters~ weeds

0N 4.7 28.0 17.4 6.6 39.5 56.6 15.7 240.6 16.4 129.7
45 kg N 4.9 25.2 21.1 5.9 40.6 52.6 16.0 271.5 45.5 119.8
ha- 1
Crimson 3.7 17.9 6.7 6.0 31.3 20.8 11.4 177.9 11.8 116.0
clover
Multivariate analysis for repeated measuresd

1989 ~ 1990 f

Sweet Lambs- Other Sweet Lambs- Other

corn quarters weeds corn quarters weeds

N source (45 kg N ha- ~ vs. crimson clover) ns ns *** *** ** *

N source × sampling date ns ns ns - ** -

aDays after plantingbCounts of lambsquarters only were taken at this date.~The substantial reduction in lambsquarters densities from first to
second count dates in all treatments was caused by an intense rainstorm at 16 DAP.dBecause of the differences in number of sampling dates
between years, emergence data for each year were analyzed separately.eBefore analysis, emergence data in 1989 were transformed in the
following manner: (y + 0.5)°5.fEmergence counts of sweet corn and other weeds in 1990, which were taken on a single date, were subjected to
univariate analysis of variance.Significance levels: ***, P < 0.001; **, P < 0.01; *, P < 0.05; ns, not significant.

s a m p l i n g d a t e ( F i g . l b ) , d i f f e r e n c e s in h e i g h t b e t w e e n at t h e first s a m p l i n g d a t e w a s 4 1 % l o w e r in t h e c l o v e r
fertilizer and clover treatments increased between the t h a n t h e f e r t i l i z e r t r e a t m e n t ( 4 . 3 vs. 7.3 l e a v e s p e r p l a n t
first a n d s e c o n d s a m p l i n g d a t e s b e f o r e d e c r e a s i n g a g a i n in 1989, 3.9 vs. 6.6 leaves per plant in 1990;
b y final h a r v e s t ( F i g . l c ) . I n b o t h y e a r s , l e a f n u m b e r P<O.O001).
 E. Dyck et al. /Agriculture, Ecosystems and Environment 56 (1995) 93-108 101

Larnbsquarters
13 a. Bi 1.8
b. S t e m ~
7
c. Height
11.25 - o m a s s / ~
1.4-
6-
oo . / / 1- 0.7
E 5-
E
0.6- c:
-- 7.75 - 0 4
4-
0.2-
6- 20406080 20 "0 60 80 20"06080
I I I I I I I 'I' i i 3 .... I ' ~ I t I
20 30 40 50 60 70 80 20 30 "0 50 60 70 80 20 30 40 50 60 70 80
Sweet Corn

12.5,,.. d. B i o m a s ~ e. Stem Diameter 7.25

2.75 - f. Heightf

c~ 10.5 - 2.25 -

f o.ts-W----- 1 6.25 - / °15V-q

_= 1.-,5- ! o.1t\ I ..q
8.5- // 0.051 k I

6.5-
2'

20
i
30
I
40
0

50
I
I
20`06080

60
'i
i i

70
I
!

80
'2'1¢ 0o:t ,
°'1
20 30
i ,
40
,
50
21'°?'°
60 70 80
5.25 1
'~

20
I
30
I
40
I
50
20 40 60 80
I
60
"t
70 80

Days after Planting

Fig. 1. Biomass, stem diameter, and height ( natural log scale ) of lambsquarters and sweet corn (averaged across weedfree and weedy treatments )
at three or four dates in 45 k g N h a - ] synthetic fertilizer ( o ) and crimson clover (e) treatments in 1989 in Experiment 1. Insets illustrate N
source × sampling date interactions, i.e. the relative difference between treatments (in I N fertilizer value] - in [crimson clover value] ) over
the sampling dates. N source × sampling date interactions for the joint analysis of 1989 and 1990 data were significant at P < 0.05 for each
growth parameter in each species.

3.1.5.3. Sweet corn growth Sweet corn biomass accumulation also manifested a
Sweet corn biomass accumulation was characterized strong N source X sampling date interaction (Table 6,
by an N source x weed X sampling date interaction Fig. ld). As with lambsquarters, the clover treatment
(Table 6). The interaction stemmed from the differ- initially negatively affected corn drymatter accumula-
ential response of fertilizer-grown and clover-grown tion; at approximately 2 weeks after emergence in 1989
sweet corn to weed presence at the last two sampling and 1990, corn biomass in the clover treatment was
dates in both years. At 50 DAP in 1989, corn biomass 25% and 38% lower, respectively, than in the fertilizer
was 33% lower in the weedy than in the weedfree N
treatment. However, at final harvest in these same
fertilizer treatment while corn grown with crimson clo-
years, biomass of clover-grown corn was 2% and 7%
ver remained unaffected by the presence of weeds. By
higher, respectively, than that of corn grown with N
final sampling date, loss of corn drymatter to weeds
was 36% in the fertilizer treatment as compared to 14% fertilizer. The greater growth of weedy corn in the clo-
in the clover treatment. In 1990, the effect of weeds ver treatment undoubtedly contributed to the decrease
was again delayed and reduced in the crimson clover in relative difference between the clover and fertilizer
treatment: biomass loss to weed interference was 19% treatments when averaged across weedy and weedfree
at 54 DAP and 21% at 86 DAP in the fertilizer treatment treatments at the last two sampling dates in either year.
as compared to no loss and a 2% loss, respectively, at However, comparison of weedfree fertilizer and clover
these same dates in the clover treatment. treatments at successive sampling periods also shows
102 E. Dyck et al. / Agriculture, Ecosystems and Environment 56 (1995) 93-108

Table 6
Aboveground biomass accumulation (g m -2) at four sampling dates in 1989 and 1990 in Experiment 1

1989

N treatment 22 DAW 36 DAP 50 DAP 77 DAP

Weedfreecorn Weedy Lambs- Weedfree Weedy Lambs- Weedfree Weedy Lambs- Weedfree Weedy Lambs-
corn quarters corn corn quarters corn corn quarters corn corn quarters

0N 0.88 0.71 1.29 12.10 10.52 18.04 58.4 33.2 101.4 239 145 294
45 kg N 1.40 1.24 1.06 12.36 18.63 18.55 95.7 64.0 108.1 305 195 325
ha-l
Crimson 0.67 1.31 0.38 14.51 9.15 9.37 74.4 89.8 61.0 274 235 203
clover

1990

N treatment 23 DAP 37 DAP 54 DAP 86 DAP

Weedfreecorn Weedy Lambs- Weedfree Weedy Lambs- Weedfree Weedy Lambs- Weedfree Weedy Lambs-
corn quarters corn corn quarters corn corn quarters corn corn quarters

0N 3.22 3.20 1.11 37.78 47.44 18.39 279.0 202.9 165.8 609 469 152
45 kg N 3.75 5.22 1.91 49.77 51.34 23.98 340.9 276.1 143.4 669 529 214
ha- J
Crinxson 2.81 2.78 0.37 52.76 55.03 7.61 288.2 321.0 63.2 646 635 124
clover
Multivariate analysis for repeated measuresb

Lambs- Sweet
quarters corn

Between effects
N source (45 kg N ha- ~vs. crimson clover) **** ns
Weed ~ - ****
N source × weed - ns

Within effects
Year * ****
N source x sampling date * **
Weed × sampling date - **
N source × weed × sampling date - t

aDays after planting.~Before analysis, data were In-transformed to reduce heterogeneity of variances among sampling dates?Weed signifies
presence or absence of lambsquarters. Significance levels: ****, P < 0.0001; **, P < 0.01; *, P <0.05; t, P < 0.10; ns, not significant.

a pattern of initially reduced growth in the clover treat-
ment followed by recovery to levels approaching or
equivalent to those in the N fertilizer treatment
(Table 6), which suggests that accelerated growth in
the clover treatment after the first sampling date con-
tributed to the diminished difference between the two
treatments.
Sweet corn stem diameter was also affected by a N
source × sampling date interaction (P < 0.0001). The
response of corn stem diameter to N source resembled
that of biomass accumulation: in both years initially
thinner corn in the legume treatment recovered to have
stem diameters that were equivalent to or thicker than
corn stem diameters in the N fertilizer treatment
(Fig. le). Corn stem diameter was also negatively
affected by weed interference in all treatments as the
experimental season progressed (data not shown,
P<O.O001).
 E. Dyck et aL /Agriculture, Ecosystems and Environment 56 (1995) 93-108
Height development in sweet corn showed a consis-
tent response to N source (P < 0.01). Although the
relative difference in height between the two treatments
tended to decrease as the growing season progressed
(Fig. l f, P < 0.05 ), clover-grown corn was shorter than
corn grown with fertilizer at all sampling dates in both
years. Corn height was increased by the presence of
weeds in all treatments (data not shown, P < 0.01), a
probable result of corn stem etiolation in response to
shading from lambsquarters within the corn row. Corn
leaf number at the first sampling date was reduced by
9% and 4%, respectively, in 1989 and 1990 in the clover
as compared to the fertilizer treatment (data not shown,
P<O.05).
3.1.5.4. Nitrogen concentration and accumulation in
lambsquarters and sweet corn
Nitrogen concentration at final harvest in both
lambsquarters and sweet corn was characterized by a
year × N source interaction (Table 7). For both weed
and crop, N concentration did not differ between N
source treatments in 1989 but was higher in the crimson
clover treatment than the fertilizer treatment in 1990.
The presence of weeds produced equivalent reductions
in sweet corn N concentration in fertilizer and clover
treatments in both years.
Results of analysis of N accumulation at final harvest
paralleled that of biomass accumulation for both lambs-
quarters and corn (Table 7). In 1989 and 1990, N accu-
mulation in clover-grown lambsquarters was 38% and
29% lower, respectively, than in lambsquarters grown
with N fertilizer. Sweet corn N accumulation were
characterized by a N source × weed interaction: in 1989
and 1990 weed interference reduced N accumulation
in corn by 47% and 35%, respectively, in the fertilizer
treatment as compared to 26% and 12%, respectively,
in the legume treatment.
3.2. Experiment 2
Biomass accumulation in the crimson clover phase
of Experiment 2 was 6267 kg h a - ~while N concentra-
tion before clover incorporation was 23.1 g kg-1. To
facilitate comparison between Experiments 1 and 2,
Experiment 2 was analyzed by contrasting the crimson
clover treatment with the fertilizer treatment of 45 kg
N h a -j.
103
Although equal applications of lambsquarters seed
had been made to all experimental units, season-long
lambsquarters density in the crimson clover treatment
was lower (P < 0.05) than in the N fertilizer treatment
(68 vs. 107 plants m - 2, respectively). Patterns in dry-
matter accumulation in Experiment 2 were consistent
with those from both years in Experiment 1, i.e. bio-
mass accumulation of lambsquarters was significantly
lower in the crimson clover plots than in the N fertilizer
plots throughout the experiment (Table 8, Fig. 2a),
remaining 34% lower at the final sampling date. Lower
biomass accumulation in the clover as compared to the
fertilizer treatment in Experiment 2 could be explained
by the difference in densities between the two treat-
ments. However, lambsquarters biomass considered on
a per plant basis was also significantly lower in the
clover than the fertilizer treatment (data not shown;
P < 0.05), which indicates that the reduced growth of
lambsquarters in the legume treatment resulted from
factors other than reduced plant density.
Lambsquarters stem diameter was negatively
affected (P < 0.10) by the clover treatment, although
the effect clearly did not persist throughout the exper-
imental season (Fig. 2b). Lambsquarters plants were
shorter in the clover than the fertilizer treatment
(P < 0.001 ). Height development in the two treatments
showed a pattern similar to that in Experiment 1; i.e.
the difference in relative heights between the two treat-
ments initially increased (from 23 to 51 DAP) before
decreasing at the end of the season (Fig. 2c). Leaf
number at first sampling date was lower (P < 0.01 ) in
the clover than the fertilizer treatment (4.1 vs. 5.1
leaves per plant, respectively).
Results of analysis of N concentration and accumu-
lation at final harvest in Experiment 2 are also in keep-
ing with Experiment 1. While N concentration in
clover-grown lambsquarters was higher than that in
lambsquarters grown with N fertilizer, N accumulation
remained lower in the clover than the fertilizer treat-
ment (Table 8).
3.3. Nitrogen source effects on crop-weed
interference
The results of Experiment 1 demonstrate a differ-
ential effect of a crimson clover N source on crop and
weed growth in comparison to that of fertilizer N.
Although the growth of both lambsquarters and sweet
104 E. Dyck et al./ Agriculture, Ecosystems and Environment 56 (1995) 93-108

Table 7
N concentration (g k g - 1) and N accumulation ( g m -2) in sweet corn and lambsquarters at final harvest dates in 1989 and 1990 in Experiment 1

N N concentration N accumulation
treatment
1989 1990 1989 1990

weedfree weedy corn lambs- weedfree weedy Iambs- weedfree weedy lambs- weedfree weedy lambs-
corn quarters corn corn quarters corn corn quarters corn corn quarters

0N 21.1 16.5 25.4 15.1 11.6 23.5 5.06 2,42 7.49 9.18 5.50 3.53
45 kg N 22.1 18.5 28.2 14.9 12.1 21.9 6.75 3.60 9.12 9.94 6.48 4.38
ha-1
Crimson 21.8 18.3 28.1 16.3 14.5 25.8 5.97 4.39 5.69 10.57 9.31 3.12
clover

Multivariate analysis for repeated measures

N concentration N accumulation

Sweet corn Lambsquarters Sweet corn Lambsquarters

Between effects
N source (45 kg N ha- l vs. crimson clover) t ns t ***
Weed a **** _ **** _

N source x weed ns - *
Within effects
Year **** **** **** ****
N source X year ** t ns ns
Weed × year ns - *
N source X weed x year ns - ns -

aWeed signifies presence or absence of lambsquarters.Significance levels: ****, P<0.0001; ***, P<0.001; **, P<0.01; *, P<0.05; j,
P < 0.10; ns, not significant

Table 8
Aboveground biomass accumulation at five sampling dates and N concentration and N accumulation at final harvest in Experiment 2

N treatment Biomass accumulation (g m -2) N concentration N accumulation

(gkg -t) ( g i n -2)
23 DAP ~ 37 DAP 51 DAP 66 DAP 79 DAP 79 DAP 79 DAP

0N 0.650 10.31 78.6 104.1 186.7 31.8 5.93

45 kg N ha- ~ 0.901 12.08 110.4 209.0 335.0 31.3 10.39
Crimson clover 0.336 5.69 47.0 119.8 222.1 34.8 7.73
Multivariate analysis for repeated measures

Biomass N concentrationc N accumulation

accumulationb

N source (45 kg N h a - ' vs. crimson clover) *** * t

N source × sampling date ns - -

aDays after planting.~Before analysis, biomass data were In-transformed to reduce heterogeneity of variance among sampling dates.ON concen-
tration and accumulation data were subjected to univariate analysis of variance. Because of recalcitrant heterogeneity of variances, N accumulation
data were rank-transformed before analysis.Significance levels: ***, P < 0.001; *, P < 0.05; t, P < 0.10; ns, not significant.
 E. Dyck et al. /Agriculture, Ecosystems and Environment 56 (1995) 93-108
13 "
a. 8 i o r n a ~ b. StemDiameter
1.3
0.8
1.2 0.6
c 0.4
0.3
5 I ! r
20 40 60 80 20 40
Days after Planting
Fig. 2. Lambsquarters biomass, stem diameter, and height (natural log scale) at four or five dates in 45 kg N ha- ~syntheticfertilizer ( o ) and
crimson clover (e) treatments in Experiment 2. Insets illustrate N source× sampling date interactions, i.e. the relative difference between
treatments (In N fertilizer value ] - In [crimsonclover valuel ) over the samplingdates. Nitrogensource× samplingdate interactions were not
significant.
corn was initially suppressed by the clover treatment,
the inhibitory effect of the clover was more severe on
lambsquarters growth than on that of sweet corn. At 2
weeks after emergence, e.g. reduction in biomass accu-
mulation (averaged over 2 years) in the clover treat-
ment was 72% for lambsquarters and 31% for sweet
corn in comparison to lambsquarters and sweet corn in
the fertilizer treatment. Reduction in growth also per-
sisted through the growing season in clover-grown
lambsquarters as compared to clover-grown sweet corn
(Fig. 1). Continued suppression of lambsquarters
growth in the clover treatment throughout the growing
season was also seen in Experiment 2, which suggests
that sweet corn interference was not responsible for the
persistence of reduced lambsquarters growth in the clo-
ver treatment in Experiment 1.
As a result of reduced interference from lambsquart-
ers, in both years of Experiment 1 use of a crimson
clover N source increased total sweet corn biomass
grown under weed-infested conditions by 20% in com-
parison to use of 45 kg N h a - 1. In fact, biomass accu-
mulation of weed-infested sweet corn grown with 180
kg N ha ~ did not exceed biomass accumulation of
weed-infested, clover-grown sweet corn in either year
(220 vs. 235 g m - 2 in 1989 and 581 vs. 635 g m -2 in
1990), a result which suggests that increased fertili-
zation is not an efficient substitute for the weed sup-
pression gained from the clover treatment.
105
6.5
c. Fleight ""
55-
4.5- 0,6""
=
0.4
35-
20 40 60 80
r 2.5
20 4O 6O 80
60 80 20
I
40 610 80
3.4. Evidence as to the causes of the inhibitory effect
of crimson clover residue
The major symptoms of the inhibitory effect of crim-
son clover in both species were suppressed growth and
development, particularly during the period between
planting and the first sampling date (approximately two
weeks after emergence). Although the experiments
described above were not primarily designed to inves-
tigate mechanisms, soil sampling results in Experiment
1 provide some evidence as to the factor(s) responsible
for the inhibitory effect of the clover treatment on early
plant development. In terms of the hypothesis of tem-
poral differences in N availability between legume and
fertilizer N sources, soil nitrate levels were initially
lower in the legume than the fertilizer treatment. Low
initial N availability could help to explain the clover's
suppression of early growth of sweet corn and lambs-
quarters, and, since nitrate is know to stimulate ger-
mination in a number of weed species, including
lambsquarters (Williams and Harper, 1965; Karssen
and Hilhorst, 1992), could also explain suppression of
weed emergence. However, soil sampling results of
Experiment 1 are not entirely consistent with the N
availability hypothesis: in 1990, initial soil nitrate lev-
els between the clover and the 0 N treatments were
equivalent even though emergence and early growth of
the lambsquarters were lower in the clover than the 0
N treatment (Tables 3, 5 and 6), a result which sug-
106 E. Dyck et al. / Agriculture, Ecosystemsand Environment 56 (1995) 93-108
gests that factors in addition to or other than low N
availability contributed to the inhibitory effect of the
clover treatment.
The existence of a mechanism other than low nitrate
availability is also suggested by the consistent lag in
height development of clover-grown lambsquarters in
Experiments 1 and 2 and the reduced height of clover-
grown sweet corn in Experiment 1. Nitrogen deficiency
results in general stunting of growth and therefore
should have similarly affected biomass accumulation
and stem diameter development. The lag in height
development may indicate inhibition of cell division
and elongation or interference with plant growth reg-
ulators, effects which have been produced by allelo-
chemicals in bioassay studies (Einhellig, 1986).
Results of plant tissue analysis for N concentration
do not provide further evidence as to the cause(s) of
the inhibitory effect of the clover treatment. Interpre-
tation of results of tissue analysis is complicated by the
fact that N concentration in plant tissue is affected by
a number of factors in addition to external N supply,
including plant growth (Glass and Siddiqi, 1984).
Because N concentration decreases with biomass accu-
mulation and because N addition stimulates growth, an
increased N supply may result in decreased plant tissue
N concentration (Greenwood et al., 1986). In this con-
text, the cause of the higher N concentrations found in
clover-grown than fertilizer-grown lambsquarters at
final harvests in Experiment 1 (in 1990) and Experi-
ment 2 remains ambiguous.
3.5. Conclusions and future research needs
The substantial reduction in weed interference with
crop growth that resulted from substitution of legume
green manure for synthetic N fertilizer in this study
demonstrates that legume green manures have the
potential to reduce the need for herbicide as well as
synthetic N fertilizer applications in subsequent crops.
However, the results of these experiments also indicate
potential drawbacks to use of a legume N source,
including possible inhibition of corn emergence (fur-
ther investigated in an experiment reported in Dyck and
Liebman, 1994). Additionally, use of crimson clover
as an N source, while substantially reducing weed
growth, did not entirely eliminate weed interference
with crop growth nor did it prevent lambsquarters
plants from setting seed by the end of the growing
season, a result that could lead to increased weed den-
sities in a cropping system in subsequent years. Clearly,
use of crimson clover needs to be considered as just
one component of a weed management strategy to be
used in conjunction with other weed-suppressing prac-
tices, including, e.g. cultivation and tillage, crop rota-
tion, and use of weed-suppressive crop varieties.
A major limitation of this study is that the effect of
N source on crop-weed interference could not be
assessed in terms of crop reproductive yield. This
resulted from the mistaken assumption that spring-
planted crimson clover could emulate the performance
of such overwintering legumes as alfalfa in developing
sufficient early season biomass accumulation to allow
early summer plowdown and subsequent successful
production of a sweet corn crop. In areas with short
growing seasons such as Maine, the effect of crimson
clover vs. fertilizer N on weed-crop interference should
be further tested in more agronomically feasible doub-
lecropping systems, e.g. crimson clover followed by a
cold-tolerant or transplanted vegetable crop. (The
effect of crimson clover on crop-weed interference in
a rotation system in which the clover is grown the year
preceding a cash crop is reported in Dyck and Liebman,
1995)
Further investigation is also needed to determine
whether the results found in this study are species spe-
cific or are more generally applicable in other cropping
systems, i.e. through the use of other legume species
(particularly those with potentially high N fertilizer
equivalency values), with other crops (including those
with growth strategies differing from that of corn, e.g.
small grains or fast-growing brassica species), and with
other weeds (including other broadleaved species as
well as annual and perennial grasses). Additional infor-
mation also needs to be generated on how specific man-
agement practices contribute or detract from the
weed-suppressive effect of crimson clover found in this
study, e.g. the effect of residue placement (surface Or
subsurface), amount of residue incorporated, depth of
incorporation, and relative timing of legume incorpo-
ration and crop planting. Finally, determination of the
mechanism(s) responsible for the weed-suppressive
effect of crimson clover in this study could lead to better
manipulation of cropping systems to maximize both N
supply and weed control from use of a legume N source.
 E. Dyck et al. /Agriculture, Ecosystems and Environment 56 (1995) 93-108
Acknowledgements
T h e a u t h o r s w o u l d like to t h a n k W . H a l t e m a n for
statistical a d v i c e a n d G. D i c k e y , G. E l d r e d , J. G o n t o s k i ,
K. G e r o w , C. Grallert, J. Kline, E. L a n g h a m , E. M e l -
lander, K. M e l l a n d e r , B. M u r p h y , K. Sader, R. Stafford,
a n d V. T r a c y for t h e i r i n v a l u a b l e h e l p w i t h the field
w o r k r e q u i r e d for this study. T h i s w o r k w a s s u p p o r t e d
by g r a n t s f r o m the J e s s i e S m i t h N o y e s F o u n d a t i o n ; the
Maine Department of Agriculture, Food, and Rural
R e s o u r c e s ; a n d the N a t i o n a l S c i e n c e F o u n d a t i o n
( D E B 9 1 2 2 8 3 3 ) a n d is c o n t r i b u t i o n 1914 o f the M a i n e
A g r i c u l t u r a l a n d F o r e s t E x p e r i m e n t Station.
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