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ReseaRch

Competitive Ability and Phytotoxic Potential of Four Winter Canola Hybrids as Affected by Nitrogen Supply
Ioannis Vasilakoglou,* Kico Dhima, Nikitas Karagiannidis, Thomas Gatsis, and Konstantinos Petrotos

aBsTRacT
Low inputs in nitrogen may affect canola (Brassica napus L.) competitiveness and allelopathic potential as well as grain and oil yields. A 2-yr study was conducted to assess nitrogen supply effect on the ability of four canola hybrids to compete with corn poppy (Papaver rhoeas L.) as well as its effect on canola seed and oil yields. Phytotoxic potential of canola hybrids on germination and growth of winter wild oat (Avena sterilis spp. ludoviciana L.), corn poppy, and wild mustard (Sinapis arvensis L.) was also determined using a perlite-based bioassay. At canola blossom, corn poppy plant number in nitrogen-treated plots was 23% less than that in the nitrogen-untreated ones. Canola seed and oil yield was reduced 18.6 and 23.7%, respectively, by the competition of 100 corn poppy plants m2, with Elan and Titan the most productive hybrids. Nitrogen supply did not increase in all cases the competitive ability or the seed and oil yields. In the laboratory, germination of the winter weeds was completely inhibited by the greatest extract concentration (5 g 100 mL1) of all hybrids. Nitrogen supply did not affect, in most cases, the phytotoxicity of canola hybrids, while phytotoxicity did not significantly differ among hybrids. Conclusively, the hybrids Elan and Titan, cultivated without any herbicide or nitrogen fertilizer, could be viable as a shortterm alternative crop system for canola seed and oil production, especially in organic or low-input fields infested by corn poppy.

I. Vasilakoglou, Dep. of Plant Production, Technological and Educational Institute of Larissa, 411 10 Larissa, Greece; K. Dhima, N. Karagiannidis, and Th. Gatsis, Dep. of Plant Production, Technological and Educational Institute of Thessaloniki, 574 00 Echedoros, Greece; K. Petrotos, Dep. of Biosystems Engineering, Technological and Educational Institute of Larissa, 411 10 Larissa, Greece. Received 20 May 2009. *Corresponding author (vasilakoglou@teilar.gr). Abbreviations: AC, ability to compete; AWC, ability to withstand competition; HB, Harper and Berkenkamp growth stage; Y, quantum yield of photosystem II.

inter canola (Brassica napus L.), or oilseed rape, has become an important economic crop in many areas worldwide, and it is usually cultivated in a crop rotation that includes winter cereals (Rathke et al., 2005; Brennan and Bolland, 2007; Hanson et al., 2008). Canola seed yield and oil content are closely related to available N (fertilizer plus soil NO3 N) ( Jackson, 2000). Compared with cereals, winter canola requires a higher amount of nutrients, and available nitrogen frequently limits seed yield. Colnenne et al. (1998) reported that canola has higher critical N demand for biomass formation than wheat. Successful weed management is essential to canola production (Martin et al., 2001), and cultivars with characteristics that provide great competitive ability (time to emergence, biomass accumulation, or plant height) might be the best choice to suppress weed growth and interference (Beckie et al., 2008). However, a wide variation in competitive ability against weeds exists among canola cultivars (Beckie et al., 2008). Canola hybrids have been introduced as a possible means of increasing yield due to heterosis (Diepenbrock, 2000) and their greater competitive ability
Published in Crop Sci. 50:111 (2010). doi: 10.2135/cropsci2009.05.0270 Published online DATE. Crop Science Society of America | 5585 Guilford Rd., Madison, WI 53711 USA
All rights reserved. No part of this periodical may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval system, without permission in writing from the publisher. Permission for printing and for reprinting the material contained herein has been obtained by the publisher.

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compared with the open-pollinated cultivars (Zand and Beckie, 2002). Allelopathy is also a major component of weed-crop interference, and it could be supplementary to integrated weed management (Kim and Shin, 2003). All species of the genus Brassica contain glycosinolates, which are not phytotoxic substances but they break down to isothiocyanates or thiocyanates (Fenwick et al., 1989). Isothiocyanates are likely to contribute to the allelopathic effects observed with decomposing Brassica tissues (Al-Khatib et al., 1997). Many isothiocyanates and thiocyanates have also been reported as strong inhibitors of seed germination and plant growth ( Ju et al., 1983; Dale, 1986; Teasdale and Taylorson, 1986; Bialy et al., 1990). Corn poppy (Papaver rhoeas L.) is an annual herbaceous plant up to 90-cm tall, with branched taproot and erect, stiffly branched hairy stems (Mitich, 2000). It is the most important dicot weed in winter cereals in Greece (Damanakis, 1983) and other areas of southern Europe that have a Mediterranean climate (Torra and Recasens, 2008). Its ability to invade, grow, and persist in cereal fields could be attributed to the formation of an abundant seed bank, an extended period of germination, and a high seed production (Holm et al., 1997). However, competition between corn poppy and winter dicot crops has not been well studied. Despite the fact that canola cultivars with great competitive and/or phytotoxic ability could be a useful tool in sustainable agriculture systems, they have not been determined sufficiently. Furthermore, data related to the role of nitrogen supply on canola competitive ability and its phytotoxicity on winter weeds are limited. The objectives of this research were (i) to assess the effect of nitrogen supply on the competitive ability against corn poppy as well as on yields of four canola hybrids in the field and (ii) to investigate inhibitory effects of these hybrids aqueous extracts on sterile oat (Avena sterilis spp. ludoviciana L.), corn poppy, and wild mustard (Sinapis arvensis L.), three of the most important winter weeds in Greece (Damanakis, 1983).

assessment made during the 20042005 growing season), one of the most important winter broadleaf weeds in Greece (Damanakis, 1983). Corn poppy density mentioned above was greater than that typically observed in Greek canola fields.

Treatments and experimental Design

Four canola hybrids (Elan, Titan, PR46w31, and PR45w04) were planted by hand in 40-cm rows to achieve an approximate density of 63 seeds m2, which reflects the common practice in Greek canola fields. Canola hybrids have recently been introduced in Greece to replace open-pollinated cultivars, and these four hybrids belong to the most cultivated ones. The planting dates were 12 Oct. 2005 and 15 Oct. 2006. Two days before canola planting, 40 kg P ha1 as superphosphate were broadcastapplied and incorporated into soil of all experimental plots. Carbofuran (2,3-dihydro-2, 2-dimethylbenzofuran-7-ylmethylcarbamate) was applied at 1.2 kg ha1 at canola planting for general insect management. In both years, the experimental area was irrigated two times with a drip irrigation system and a total volume of 60 mm of water. In Year 1, irrigation was applied during May, whereas the experimental area was irrigated during April in Year 2. This was done because of the low rainfall recorded during May in Year 1 and April in Year 2 (Fig. 1). The previous crop was two-row barley (Hordeum vulgare L.), which was harvested in mid-June 2005. Barley straw was baled and removed after harvest, and barley volunteer plants were observed at very low densities during the experiment. Other common cultural practices were imposed as needed during the growing season. A split-split-plot arrangement of treatments was employed in a randomized complete block design with four replicates. Main plots consisted of two nitrogen fertilizer levels (without N supply and with 100 kg N ha1 as urea 2 d before planting) with a plot size of 30 5 m. Urea was broadcast-applied and incorporated into the soil, reflecting the common practice in Greek canola fields. All main plots were separated by a 3-m wide alley. Each main plot was divided into four 6 5 m subplots of 12 canola rows each. The four canola hybrids were the subplots. All subplots were separated by a 2-m wide alley. Each subplot was subdivided into two 2.5 5 m areas of six canola rows each, one that remained weed free and one that was infected by corn poppy. All sub-subplots were separated by a 1-m wide alley and were left weedy until 15 December, in order for complete corn poppy emergence. At that time, competition had not started, as canola and corn poppy were at their early growth stages (four- to six-leaf and cotyledon- to four-leaf stage, respectively) (McMullan et al., 1994; Yaghoobi and Siyami, 2008). However, on 15December of each year, corn poppy was hand removed in half of all sub-subplots (weed-free sub-subplots). In weedy sub-subplots, corn poppy uniformly emerged; however, it was thinned, as needed, to a density of about 100 plants m2 and allowed to grow in competition with canola until harvest.

MaTeRIaLs aND MeThODs

Field experiment
Experimental Site
A field experiment was conducted in 20052006 (Year 1) and was repeated in 20062007 (Year 2) at the Technological and Educational Institute Farm of Thessaloniki in northern Greece (224410 E, 403706N). Experiments were established on a sandy loam (Typic Xeropsamment) soil whose physicochemical characteristics were sand 644 g kg1, silt 280 g kg1, clay 76 g kg1, organic matter content 10 g kg1, and pH (1:2 H2O) 7.6. Preplant soil analysis conducted in the middle of September showed that initial nitrate content ranged from 84 to 88 and 89 to 93 mg kg1 of soil in Year 1 and Year 2, respectively. Mean monthly temperature and rainfall data recorded near the experimental area are shown in Fig. 1. The experimental area was naturally infested by 110 to 120 plants m2 of corn poppy (as confirmed by a visual 2

Data Collection
The canola stand was counted at 6 wk after planting in the two central canola rows of each sub-subplot. Simultaneously, corn poppy density was measured in a 5 0.8 m area between the two central interrows of each sub-subplot. Corn poppy plant number and fresh weight of both species were determined in a 5 0.4 m area at the middle of the blossom stage of canola at 27 wk after
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Fig. 1. Total monthly rainfall and mean monthly temperature during the experiment.

Bioassay Procedure
Ten canola plants of each hybrid studied in the field experiment were randomly harvested from the weed-free sub-subplots at the middle of blossom growth stage (HB 4.24.3) (Harper and Berkenkamp, 1975) in each year. The harvested plants were chopped into 5-cm-long pieces, dried at room temperature (25C) for 72 h, and ground in a Wiley mill (Thomas Scientific, Swedesboro, NJ) through a 1-mm screen. Then, aqueous extracts (w/v) were prepared in 400-mL glass jars by adding 1.25, 2.5, 5.0, or 10.0 g from each plant sample to 200 mL deionized water and shaking in a horizontal shaker for 4 h at 200 rpm. Solutions were filtered through four layers of cheesecloth to remove fiber debris, centrifuged at 1750 g in a centrifuge with a 30-cm rotor diameter for 1 h, and supernatants were then filtered through a layer of filter paper (Whatman No. 42; W. & R. Balston, Maidstone, Kent, UK). Extracts were stored for 2 to 6 d at 4C until bioassayed. There were three replicate extracts for each canola hybrid by extract concentration treatment (0.63, 1.25, 2.50, and 5.00 g 100 mL 1). Petri dish bioassays were performed to assess the phytotoxic effects of the four canola hybrids on winter wild oat, corn poppy, and wild mustard. In particular, germination, root length, and fresh weight of winter wild oat and wild mustard in perlite treated with each of the canola hybrid aqueous extracts were investigated. For corn poppy, only germination was evaluated because of the small size of its seedlings. Winter wild oat, corn poppy, and wild mustard seeds (40, 200, and 50, respectively) were placed in 8.5-cm diameter plastic petri dishes and were covered with 5 g of perlite. Open petri dishes were moistened with 15 mL of canola extract per petri dish from each of the canola hybrid extracts. Deionized water was used in control petri dishes. There were two petri dishes for each replicate extract, and petri dishes were arranged in a completely randomized design. Afterward, petri dishes were stored on shallow trays and placed inside a plastic bag to retain moisture. Trays were then placed in an illuminated (8 h light:16 h dark) growth chamber at 20 2C for 14 d. At the end of the incubation period, plants were removed from petri dishes, carefully washed free of perlite, and average (mean of the two petri dishes used for each replicate extract) germination as well as root length and total fresh weight (of the germinated seeds only) were measured. Measured data were expressed as a percentage of the water control. The experimental procedure was conducted twice using the canola samples taken in Year 1 and Year 2. Fungal contamination was not observed during these experiments. 3

Competition Indices
Competition indices of canola were calculated according to Watson et al. (2006). Ability to withstand competition (AWC) was calculated as AWC = 100 (Ywp/Ywfp), where Ywp is the seed yield from the weedy sub-subplot and Ywfp is the seed yield from the weed-free sub-subplot. Ability to compete (AC) was calculated as AC = 100 [(bw/bt) 100], where bw is the fresh weight of corn poppy and bt is the total fresh weight (canola and corn poppy).

Laboratory experiment
Plant Material
Seeds from mature plants of winter wild oat, corn poppy, and wild mustard were collected near the experimental area during June 2004. All collected seeds were dried in the greenhouse, air-cleaned to remove unviable seeds and other plant residues, and stored in a refrigerator at 3 to 6C until used for the experiment. The viability of winter wild oat, corn poppy, and wild mustard seeds was evaluated before use and was approximately 25, 10, and 22%, respectively.
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planting (Harper and Berkenkamp growth stage [HB] 4.24.3) (Harper and Berkenkamp, 1975). During this sampling, one (the fifth) of the six canola rows of each sub-subplot as well as the corn poppy plants found among and around this row (0.40 5 m area) of each weedy sub-subplot were clipped at the surface, and corn poppy plant density and fresh weight of both species were measured. This stage was chosen because corn poppy had the greatest biomass and the critical period of competition between canola and weed had been completed (Martin et al., 2001). Chlorophyll fluorescence and quantum yield of photosystem II (Y) were used to assess the reaction of the canola hybrids to nitrogen supply and corn poppy competition. Measurements of the chlorophyll fluorescence parameters were made at two canola growth stages (when lower buds were present but enclosed by leaves [HB 3.1] and when all viable buds on raceme were open [HB 4.9]) (Harper and Berkenkamp, 1975) using a chlorophyll fluorometer (MINI-PAM, Miniaturised Pulse-Amplitude-Modulated photosynthesis yield analyzer, Walz Company, Effeltrich, Germany) with measurement light intensity of 0.15 mol m2s1, a frequency of 0.6 kHz, and a saturation pulse intensity of 16,000 mol m2s1 for 0.8 s. Two measurements per plant were made on the upper leaves of five marked plants in the center of each sub-subplot to determine fluorescence at steady state (Fs) and the maximum fluorescence after saturation flash (Fm). Quantum yield of photosystem II was calculated using the equation developed by Genty et al. (1989): Y = (Fm Fs)/Fm. The average of the 10 measurements per sub-subplot was used for further data analysis. At harvest, canola seed yield and 1000-seed weight were determined by hand harvesting the pods of canola plants in two 5-m-long rows (the second and third) of each sub-subplot at 20June of both growing seasons. Furthermore, seed yield samples were forced-air dried at 29C to a uniform moisture level, cleaned, weighed, and a subsample of 200 g taken for seed oil content. Oil concentration was determined using the crude fat method (Association of Official Analytical Chemists, 1990). Oil yield was calculated by multiplying seed yields by the concentration of oil in seeds, both corrected to 8% moisture content.

Table 1. Significance levels for corn poppy (Papaver rhoeas L.) plant number or fresh weight data (27 wk after planting) as well as ability to withstand competition (AWC) and ability to compete (AC) data of four winter canola (Brassica napus L.) hybrids in 20052006 (Year 1) and 20062007 (Year 2).
Source
Years (Y) Replicates (Y) Nitrogen (N) YN Error Canola hybrid (CH) Y CH N CH Y N CH Error CV, %

df
1 6 1 1 6 3 3 3 3 36

Significance of F ratio Plant number Fresh weight AWC AC

* ns * ns ** ns ns ns 11.8 ns ns ns ns * ns ns ns 8.1 *** * ** * ns * ns ns ns ns ns * ns ns ns ns

variables (y), while canola aqueous extract concentration (g dry weight 100 mL 1) was the independent variable (x). Also, the phytotoxic doseresponse effects of canola extracts on winter weeds germination, root length, and fresh weight were assessed by the Whole-range assessment method (An et al., 2005). Inhibition index was calculated by Eq. [1] used by Liu et al. (2007). I = DcDn[R(0) f(D)]dD/0Dn R(0)dD [1] In this equation, concentrations tested ranged from 0 to D n, Dc was the threshold dose at which response equaled the value of control and above which the responses were inhibitory, R(0) was the response at 0 g dry matter per 100 mL (control), and f(D) represented the response function. Germination, root length, and total fresh weight inhibition areas across the whole range of canola hybrids extract concentrations and the corresponding Inhibition indices (I) were calculated separately for each replicate using the WESIA (Whole-Range Evaluation of the Strength of Inhibition in Allelopathic-Bioassay) software (Liu et al., 2007) and afterward were subjected to a combined over repetition time ANOVA. The programs SPSS (SPSS, 1997) and MSTAT (MSTATC, 1988) were used to conduct regression analyses and analyses of variance, respectively. Fishers protected LSD procedures were used to detect and separate mean treatment differences at P = 0.05.

Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.

18.0 15.5

*Significant at the 0.05 level. **Significant at the 0.01 level. ***Significant at the 0.001 level. Corn poppy plant number and fresh weight data before the ANOVA were sqrt(x) or log(x) transformed, respectively.

ns, not significant.

statistical analyses
Data received from the field experiment were analyzed over the growing season (year). A split-plot factorial approach (nitrogen supply canola hybrid) was performed for canola and corn poppy emergence (6 wk after planting), corn poppy plant number, and fresh weight of both species (27 wk after planting) as well as for AWC and AC data. Corn poppy fresh weight (bw for AC) and canola seed yield values (Ywp for AWC) measured in individual weedy sub-subplots were used as numerators. However, treatment means for canola fresh weight plus corn poppy fresh weight mean values from weedy sub-subplots (bt for AC) and canola yield from weed-free sub-subplots (Ywfp for AWC) were used as denominators to improve normality and homogeneity of data variances. Corn poppy plant number and fresh weight data before the ANOVA were square-root(x)- and log(x)-transformed, respectively, to reduce their heterogeneity, but means presented are back-transformed values. Canola data (quantum yield of photosystem II [Y], fresh weight [27 wk after planting], seed yield, oil yield, and 1000-seed weight) were analyzed by using a split-split-plot factorial approach (nitrogen supply canola hybrid corn poppy competition). Winter weed bioassay data were analyzed over repetition time using a factorial approach (nitrogen supply canola hybrid aqueous extract concentration). Data for the germination, root length, and total fresh weight of the three weed species before the ANOVA were log(x + 1) transformed to reduce their heterogeneity, but means presented are back-transformed values. Linear, quadratic, hyperbolic, exponential, and logarithmic equations were tested for their suitability to describe the relationship between weed germination, root length, and total fresh weight response and canola aqueous extract concentration. The equation with the highest adjusted coefficient of determination (R a 2) and F values was judged to be the most appropriate. In these regression equations, germination, root length, and total fresh weight (% of control) were the dependent 4

ResULTs aND DIscUssION

Field experiment
Initial nitrate content, which was greater than that typically observed during the growing season in Greek fields, could be the result of the very low rainfall recorded during summer (Fig. 1) and, consequently, of the restricted leaching of nitrate form during this period. However, only part of this nitrogen amount could be used by canola, mainly because of nitrate leaching and denitrification during winter (Colnenne et al., 2002; Sieling et al., 2006). Weed Response Corn poppy emergence at 6 wk after planting was not significantly affected by year, nitrogen supply, and canola hybrid. In particular, corn poppy density averaged 104 plants m2 (data not shown). Similarly, Beckie et al. (2008) found that open-pollinated and hybrid canola had no significant effect on total weed seedling density at 4 wk after crop emergence. However, at crop blossom (27 wk after planting), differential corn poppy suppressive ability among canola hybrids was evident, agreeing with Beckie et al. (2008). In particular, corn poppy plant number was affected by year, nitrogen supply, and canola hybrid, whereas corn poppy fresh weight was affected only by canola hybrid (Table 1). So, the means presented in Table2 are averaged across year and nitrogen supply, while the nitrogen supply effect on corn poppy plant number is discussed. At 27 wk after planting, corn poppy plant number in nitrogen-treated plots was 23% less (89 plants m2) than that in nitrogen-untreated ones (116 plants m2). The lower corn poppy plant number in nitrogen-treated
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Corn poppy plant number or fresh weight data before the analysis of variance were sqrt(x) or log(x) transformed, respectively, but the mean values presented are back transformed. Means within each column followed by the same letter are not significantly different according to the Fishers protected LSD test at P = 0.05.

Canola Response Canola emergence at 6 wk after planting was not significantly affected by year, nitrogen supply, and canola hybrid. In particular, canola density averaged 60 plants m2 (data not shown). The AWC of canola was affected by year and nitrogen supply, as well as by year nitrogen supply and year canola hybrid interactions (Table 1). However, the AC of canola was affected only by year nitrogen supply interaction. In Year 1, AWC did not differ between nitrogen supply treatments and among canola hybrids (ranged from 75.9 to 78.4%). However, in Year 2, AWC in nitrogen-treated plots was greater than that in nitrogen-untreated ones (97.1 and 85.8%, respectively). Safahani Langeroudi and Kamkar (2009) found that winter canola AWC of wild mustard ranged from 4 to 47%. In Year 1, AC was greater in nitrogen-treated plots, as compared with the nitrogen-untreated ones (81.2 and 65.1%, respectively), whereas it was not affected by nitrogen supply in Year 2 (ranged from 72.2 to 75.5%). The lack of difference between AC values in nitrogen-treated and nitrogen-untreated plots in Year 2 could be attributed to low rainfall recorded during January to April (Fig. 1), which may have resulted in lower nitrogen utilization efficiency and lower canola early growth rate, as compared with Year 1. The greatest AWC value was provided by the hybrid Elan. Generally, nitrogen supply did not improve the competitive ability of all canola hybrids and for each hybrid in both years, perhaps because of the satisfactory initial nitrate content in soil and the different nitrogen utilization efficiency among hybrids. At HB 3.1 (before blossom) canola growth stage, quantum yield of photosystem II (Y) was not affected by year, nitrogen supply, canola hybrid, and corn poppy competition. However, at HB 4.9 (end of blossom) canola growth stage, the Y was affected by year (P < 0.01), canola hybrid (P < 0.01), and corn poppy competition (P < 0.001) as well as by year canola hybrid (P < 0.01) and year corn poppy competition (P < 0.01) interactions. The year canola hybrid and year corn poppy competition interaction means are presented in Fig. 2. In Year 1, canola Y was 10% lower than that in Year 2 (Fig. 2A). The greater rainfall recorded
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during May (period of blossom) in Year 2 as compared with Year 1 (Fig. 1) could account for the greater Y of the canola hybrids as a result of the greater nitrogen utilization efficiency and the greater radiation use efficacy (Ogunlela et al., 1989; Diepenbrock, 2000; Colnenne et al., 2002). The hybrids Elan and Titan provided greater Y than the other two hybrids in Year 1, whereas canola Y was not affected by hybrid in Year 2. Generally, the corn poppy competition caused the decrease of canola Y (Fig. 2B). The fact that the nitrogen supply did not affect the canola Y could be attributed to the satisfactory nitrogen amount in soil before canola planting, associated with the results reported by Ogunlela et al. (1989), who found, in a greenhouse experiment, that

Fig. 2. Quantum yield of photosystem II at the end of blossom growth stage of four canola (Brassica napus L.) hybrids as affected by (A) year canola hybrid and (B) year corn poppy competition interactions. Means within each interaction followed by the same letter are not significantly different according to the Fishers protected LSD test at P = 0.05.

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plots, as compared with that in nitrogen-untreated ones, could be attributed to the greater soil nitrogen concentration in nitrogen-treated plots, which, associated with the extended canola root system, should result in greater nitrogen use by canola hybrids and, consequently, in greater early canola growth (Colnenne et al., 2002). The lower corn poppy plant number and fresh weight recorded in the hybrid PR46w31, as compared with those recorded in the hybrids Elan and PR45w04 (Table 2), could be attributed to greater nitrogen use efficiency at vegetative stage, resulting in different growth rates among canola hybrids (Svecnjak and Rengel, 2006).

Table 2. Plant number and fresh weight of corn poppy (Papaver rhoeas L.) grown in four canola (Brassica napus L.) hybrids at the middle of blossom growth stage (27 wk after planting). Means are averaged across year and nitrogen supply.
Hybrids
Elan Titan PR46w31 PR45w04

Plant number
m 2 118 a 93 ab 89 b 110 a

Fresh weight
kg m2 2.26 a 1.51 bc 1.16 c 2.04 ab

Fig. 3. Seed yield of canola (Brassica napus L.) as affected by (A) year canola hybrid corn poppy competition and (B) nitrogen supply canola hybrid corn poppy competition interactions. Means within each interaction followed by the same letter are not significantly different according to the Fishers protected LSD test at P = 0.05.

the increase in N supply from 100 to 170 mg L1 did not significantly affect the content of chlorophyll a and b or the chlorophyll a/b ratio in canola leaves. Canola fresh weight at 27 wk after planting was affected by year (P < 0.001), nitrogen supply (P < 0.05), corn poppy competition (P < 0.001), and year corn poppy competition interaction (P < 0.001). Generally, the fresh weight of canola plants receiving the nitrogen supply was 22% greater than that of plants in nitrogen-untreated plots (7.53 and 6.15 kg m2, respectively). As mentioned above, the greater soil nitrogen concentration in nitrogen-treated plots, associated with the extended canola root system, should result in greater nitrogen use by canola cultivars and, consequently, in greater canola growth (Colnenne et al., 2002). Similarly, Asare and Scarisbrick (1995) found that, in most cases, the nitrogen supply increased the dry matter yield of canola. In both weedy and weed-free treatments, canola fresh weight in Year 2 (7.41 and 10.26 kg m2, respectively) was greater than in Year 1 (4.35 and 5.35 kg m2, respectively). The greater temperatures recorded during January to April in Year 2 as compared with Year 1 (Fig. 1) could account for this difference in canola fresh weight as long as canola water needs had been satisfied by rainfall or irrigation. In both years, canola fresh weight in weed-free treatments was greater than in weedy ones, mainly owing to the lack of corn poppy competition. Canola seed yield was affected by year (P < 0.01), nitrogen supply (P < 0.05), and corn poppy competition
6

(P < 0.001) as well as by year nitrogen supply (P < 0.01), year canola hybrid corn poppy competition (P < 0.05), and nitrogen supply canola hybrid corn poppy competition (P < 0.01) interactions. The year canola hybrid corn poppy competition and nitrogen supply canola hybrid corn poppy competition interaction means are presented in Fig. 3. In Year 2, canola seed yield was greater than in Year 1 in both weedy and weed-free treatments (Fig. 3A), agreeing with the fresh weight data mentioned above. In weedy plots, canola seed yield was lower than in weed-free ones by 16.5 to 21.8% (averaged across years) owing to corn poppy competition. Blackshaw et al. (1987) found that seed yield reduction of spring canola ranged from 19 to 77% with the competition of 20 to 80 wild mustard plants m2 and only 20 to 25% with the competition of common lambsquarters (Chenopodium album L.). However, canola seed yield was not affected by a stable trend in the nitrogen supply or the hybrid (Fig. 3B). In particular, in weedy conditions, nitrogen supply increased seed yield of hybrids Titan and PR45w04. In weed-free conditions, nitrogen supply increased seed yield of hybrids Elan and PR45w04. Asare and Scarisbrick (1995) found that the increase of nitrogen supply from 120 to 240 kg ha1 increased canola seed yield, mainly because of the greater number of pods on the terminal raceme and heavier seed weight. Similarly, Jackson (2000) found that increasing the nitrogen rate from 0 to 252 kg ha1 increased canola seed yield. However, Colnenne et al. (2002) did not find any difference in seed yield of canola supplied with 0 or 100 kg nitrogen ha1. According to the same researchers, this was because of satisfactory nitrogen concentration, which was released by natural mineralization in soil, allowing sufficient canola growth in autumn even though no fertilizer was applied. Canola oil yield was affected by canola hybrid (P<0.01) and corn poppy competition (P < 0.001) as well as by year nitrogen supply canola hybrid (P < 0.05), year nitrogen supply corn poppy competition (P <0.01), and nitrogen supply canola hybrid corn poppy competition (P< 0.01) interactions. The year nitrogen supply corn poppy competition and nitrogen supply canola hybrid corn poppy competition interaction means are presented in Fig. 4. In Year 1, oil content in nitrogen-untreated plots was 42.5 and 40.2% (as averaged across hybrids) in weedy and weed-free treatments, respectively (Fig. 4A). The corresponding oil content in nitrogen-treated plots was 34.2 and 36.3%, agreeing with the results reported by Asare and Scarisbrick (1995), who found that the increase in nitrogen rate decreased canola oil content because of increased seed protein content. In Year 2, oil content in nitrogenuntreated plots was 28.1 and 29.7% in weedy and weedfree treatments, respectively, whereas the corresponding oil content in nitrogen-treated plots was 29.4 and 31.0%. Canola oil yield did not differ between years, except for the
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Laboratory experiment
Germination, root length, and total fresh weight of winter wild oat, wild mustard, and corn poppy were, in most cases, significantly affected by year (P < 0.001), nitrogen supply (P < 0.001), canola hybrid (P < 0.001), aqueous extract concentration (P < 0.001), and their interaction (P< 0.001). The year nitrogen supply canola hybrid aqueous extract concentration interaction means are presented in Fig. 5, 6, and 7. In most cases, winter wild oat and wild mustard germination was affected less than root length and total fresh weight (Fig. 5 and 6). However, germination of the three winter weeds was completely inhibited by the greatest aqueous extract concentration (5 g 100 mL 1) of all canola hybrids (Fig. 5, 6, and 7). Winter weeds germination, root length, and total fresh weight decreased as canola aqueous extract concentration increased (Fig. 5, 6, and 7), agreeing with results reported by Teasdale and Taylorson (1986), who found that inhibition of large crabgrass [Digitaria sanguinalis (L.) Scop.] germination increased as the concentration of isothiocyanate increased. Quadratic regression equation (y = a + bx + cx 2) was in most cases the best fit for winter wild oat and wild mustard germination, root length, and total fresh weight changes vs. canola aqueous extract concentration (Fig. 5 and 6). However, the exponential regression equation (y = a bx) was in most cases the best fit for corn poppy germination changes vs. canola aqueous extract concentration (Fig. 7). Estimated slopes (b parameters) for germination, root length, and total fresh weight lines indicated that in most cases weed parameters reduction was greater in Year 2 than in Year 1. This difference could be attributed to greater amounts of allelochemicals produced by canola, as a result of the previously reported greater photosynthetic ability in Year 2.

Fig. 4. Canola (Brassica napus L.) oil content as affected by (A) year nitrogen supply corn poppy competition, and oil yield as affected by (B) year nitrogen supply corn poppy competition and (C) nitrogen supply canola hybrid corn poppy competition interactions. Means within each interaction followed by the same letter are not significantly different according to the Fishers protected LSD test at P = 0.05.

Winter wild oat germination, root length, and total fresh weight inhibition caused by the canola aqueous extracts in Year 1 ranged from 61.5 to 72.4, 63.9 to 85.7, and 59.9 to 74.6%, respectively (Table 3). The corresponding inhibition caused by aqueous extracts in Year 2 was 44.1 to 84.5, 79.9 to 88.5, and 69.2 to 89.1%. Nitrogen supply did not affect, in most cases, the phytotoxicity of canola hybrids in Year 1, possibly because of the satisfactory initial nitrogen content in soil. However, nitrogen supply in Year 2 resulted in greater canola phytotoxicity on germination and total fresh weight of winter wild oat. Protein content generally increases when the rate of nitrogen application is increased (Asare and Scarisbrick, 1995). The last probably leads to greater concentration of phytotoxic substances, and it could be considered a factor for the greater phytotoxicity of canola extracts. The greatest inhibitory effect on winter wild oat in Year 1, as averaged across germination, root length, and total fresh weight
7

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weed-free treatments supplied with nitrogen, when canola oil yield was greater in Year 1 than in Year 2 (Fig. 4B). Averaged across years, canola oil yield decreased 23.7% with corn poppy competition, agreeing with the abovementioned seed yield results. In nitrogen-treated and weedfree plots, the hybrids Elan and Titan provided slightly greater oil yield than the hybrids PR46w31 and PR45w04 (Fig. 4C). In corresponding weedy treatments, the cultivar Titan provided the greatest oil yield. In nitrogen-untreated and weed-free plots, the cultivar Titan provided again the greatest oil yield, while the hybrid PR45w04 provided the lowest. In corresponding weedy treatments, the greatest oil yield was provided by the hybrid Elan. The 1000-seed weight of canola was affected only by year (P < 0.001) and hybrid (P < 0.001). In particular, the hybrids PR46w31 and Titan provided greater 1000-seed weight (3.00 and 2.89 g, respectively) than the hybrids PR45w04 and Elan (2.62 and 2.56 g, respectively).

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Fig. 5. Inhibitory effect (% of water control) of four winter canola (Brassica napus L.) aqueous extracts on germination, root length, and total fresh weight of winter wild oat (Avena sterilis spp. ludoviciana L.) as affected by concentration and nitrogen supply. Lines describe quadratic regression equations [data before the ANOVA were log(x + 1) transformed, but the mean values used were back transformed]. N, without nitrogen supply; +N, with nitrogen supply; Ra2, adjusted coefficients of determination; *, **, ***, significance of F values at the 0.05, 0.01, and 0.001 probability levels, respectively; ns, not significant.

Fig. 6. Inhibitory effect (% of water control) of four winter canola (Brassica napus L.) aqueous extracts on germination, root length, and total fresh weight of wild mustard (Sinapis arvensis L.) as affected by concentration and nitrogen supply. Lines describe quadratic regression equations [data before the ANOVA were log(x + 1) transformed, but the mean values used were back transformed]. N, without nitrogen supply; +N, with nitrogen supply; Ra2, adjusted coefficients of determination; *, **, ***, significance of F values at the 0.05, 0.01, and 0.001 probability levels, respectively; ns, not significant.

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inhibition, was caused by the hybrid PR45w04 (74.0%), while the greatest inhibitory effect on winter wild oat in Year 2 was caused by the hybrid PR46w31 (80.7%). In Year 1, wild mustard germination, root length, and total fresh weight inhibition caused by the canola aqueous extracts ranged from 58.7 to 73.4, 64.3 to 74.2, and 54.3 to 78.5%, respectively (Table 4). The corresponding inhibition caused by aqueous extracts in Year 2 was 73.2 to 83.8, 72.1 to 84.6, and 74.3 to 85.2%. In both years, nitrogen supply did not affect, in most cases, the phytotoxicity of canola hybrids. The greatest inhibitory effect on wild mustard in Year 1, as averaged across germination, root length, and total fresh weight inhibition, was caused by the hybrid PR45w04 (74.4%). The greatest inhibitory effect on wild mustard in Year 2 was caused by the hybrid PR46w31 (81.3%) followed by the hybrid Titan (81.1%). Corn poppy germination inhibition caused by the canola aqueous extracts in Year 1 ranged from 65.7 to 84.0% (Table 4). The corresponding inhibition caused by aqueous extracts in Year 2 was 41.9 to 85.7%. Nitrogen supply did not affect, in most cases, the phytotoxicity of canola hybrids in Year 1. However, nitrogen supply in Year 2 resulted in greater canola phytotoxicity on corn poppy germination. The greatest inhibitory effect on corn poppy in Year 1 was caused by the hybrid Elan (81.9%) followed by the hybrid Titan (80.3%), whereas the greatest inhibitory effect in Year 2 was caused by the hybrid PR46w31 (78.8%) followed by the hybrid PR45w04 (77.1%). The recorded differences among the four canola hybrids tested for their inhibitory effect on germination, root length, and total fresh weight inhibition of the three winter weeds could be attributed to their differences in kind, total amount, and physicochemical characteristics of the allelochemicals produced. Regarding the chemistry of substances involved in allelopathy, Fenwick et al. (1989) reported that all species of the genus Brassica contain glycosinolates (not phytotoxic substances) that break down to isothiocyanates or thiocyanates. Many isothiocyanates and thiocyanates have been reported to be strong inhibitors of seed germination and plant growth ( Ju et al., 1983; Dale, 1986; Teasdale and Taylorson, 1986; Bialy et al., 1990; AlKhatib et al., 1997). In particular, allyl, methyl, benzyl, and b-phenylethyl isothiocyanates were potent inhibitors of seed germination (Teasdale and Taylorson, 1986; Dale, 1986), whereas ionic thiocyanate inhibited root or shoot growth of cabbage (Brassica oleracea L.), bean (Phaseolus vulgaris L.), and tobacco (Nicotiana tabacum L.) ( Ju et al., 1983). Also, Al-Khatib et al. (1997) found, in a closed containers experiment, that the methyl, phenyl, ethyl, and allyl isothiocyanates inhibited germination and growth of barnyardgrass [Echinochloa crus-galli (L.) P. Beauv.], redroot pigweed (Amaranthus retroflexus L.), and green pea (Pisum sativum L.) more than benzyl, butyl, propyl, and b-phenylethyl isothiocyanates did.
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Fig. 7. Inhibitory effect (% of water control) of four winter canola (Brassica napus L.) aqueous extracts on germination of corn poppy as affected by concentration and nitrogen supply. Lines describe exponential regression equations [data before the ANOVA were log(x + 1) transformed, but the mean values used were back transformed]. N, without nitrogen supply; +N, with nitrogen supply; Ra2, adjusted coefficients of determination; *, **, ***, significance of F values at the 0.05, 0.01, and 0.001 probability levels, respectively; ns, not significant.

Table 3. Growth inhibition of winter wild oat (Avena sterilis spp. ludoviciana L.) caused by aqueous extracts of four canola (Brassica napus L.) hybrids as assessed by the inhibition indices estimated from the use of the WESIA (Whole-Range Evaluation of the Strength of Inhibition in Allelopathic-Bioassay) software.
Germination Hybrid + Winter wild oat Root length Total fresh weight Nitrogen supply + +

Elan Titan PR46w31 PR45w04 Elan Titan PR46w31 PR45w04

Inhibition index (%) Year 1 64.4 fgh 63.7 fgh 85.7 a 79.9 a 72.4 cd 69.6 de 84.0 a 63.9 b 61.5 h 67.4 ef 82.8 a 82.3 a 67.0 efg 71.0 cde 82.5 a 83.8 a Year 2 44.4 i 70.1 cde 82.4 a 82.9 a 44.1 i 78.9 b 79.9 a 88.5 a 62.8 gh 84.5 a 84.4 a 87.3 a 70.8 cde 74.6 bc 85.5 a 83.4 a

66.8 fg 74.6 cde 64.3 gh 72.0 def 69.2 fg 67.4 fg 75.8 cde 77.4 c

59.9 h 67.5 fg 70.6 ef 67.8 fg 82.8 b 85.9 ab 89.1 a 77.2 cd

Means within germination, root length, or total fresh weight columns followed by the same letter are not significantly different according to the Fishers protected LSD test at P = 0.05.

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Table 4. Growth inhibition of wild mustard (Sinapis arvensis L.) or corn poppy (Papaver rhoeas L.) caused by aqueous extracts of four canola (Brassica napus L.) hybrids as assessed by the inhibition indices estimated from the use of the WESIA (WholeRange Evaluation of the Strength of Inhibition in Allelopathic-Bioassay) software.
Germination Hybrid
Year 1 Elan Titan PR46w31 PR45w04 Year 2 Elan Titan PR46w31 PR45w04

Wild mustard Root length +

Total fresh weight Nitrogen supply + +

Corn poppy Germination +

Inhibition Index (%)

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61.3 ef 63.6 ef 66.8 de 73.4 cd 73.2 cd 78.4 abc 76.6 c 83.5 ab

63.3 ef 58.7 f 67.4 de 73.3 cd 75.9 c 83.8 a 77.8 abc 76.9 bc

64.3 f 67.7 ef 71.7 de 72.8 de 72.1 de 78.5 bc 79.3 abc 84.6 a

64.5 f 67.7 ef 72.8 de 74.2 cd 78.8 bc 80.2 ab 79.7 ab 79.8 ab

65.7 f 68.0 ef 76.2 cd 73.9 cde 74.3 cde 80.5 abc 77.8 c 84.8 ab

70.0 def 54.3 g 77.9 c 78.5 bc 78.7 abc 85.2 a 78.9 abc 78.4 bc

78.8 bcde 84.0 ab 75.7 def 65.7 h 41.9 j 59.6 i 71.9 fg 77.2 cdef

81.8 abc 79.8 bcd 73.3 ef 72.4 fg 66.6 gh 79.9 bcd 85.7 a 77.0 cdef

Means within germination, root length, or total fresh weight columns followed by the same letter are not significantly different according to the Fishers protected LSD test at P = 0.05.

cONcLUsIONs
The results of this study indicated that nitrogen and herbicide usage is essential to maximize canola yield. However, canola hybrids Elan and Titan cultivated without any herbicide usage or nitrogen fertilizer could be viable as short-term alternative crop systems for canola seed and oil production, especially in organic and low-input system fields infested by corn poppy, but with satisfactory initial nitrogen content. acknowledgments
Authors are grateful to Dr. Christo Dorda for providing canola hybrids. This research was funded by the Research Committee of the Technological and Educational Institute of Thessaloniki.

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