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Weed Biology and Management 12, 1221 (2012)

RESEARCH PAPER

Inuences of long-term different types of fertilization on weed community biodiversity in rice paddy elds
wbm_430 12..21

KAIYUAN WAN1, YONG TAO1, RUHAI LI2, JUNFENG PAN1, LEILEI TANG1,3 and FANG CHEN1* Key Laboratory of Aquatic Botany and Watershed Ecology, Wuhan Botanical Garden, Chinese Academy of Sciences, 2 Institute of Plant Protection and Soil Science, Hubei Academy of Agriculture Sciences, Wuhan and 3 Life Science College, Graduate University of the Chinese Academy of Sciences, Beijing, China
In order to provide a scientic basis for developing integrated weed management strategies in rice paddy elds, this study investigated the inuences of different types of fertilization on weed biodiversity.The experiment was conducted at Long-term-located Monitoring Station for Soil Fertility,Agricultural Science Academy, of Jiangxi Province, China. Five fertilization treatments were set: no fertilization (NOF), PK, NP, NK, and NPK.The results showed that the inuence of different fertilization treatments on weed community traits followed the models PK > NOF > NK > NP > NPK for species richness, PK > NOF > NK > NP > NPK for species diversity, NPK > NP > NK > NOF > PK for community dominance, and PK > NOF > NK > NP > NPK for community evenness. Under NPK (i.e. balanced fertilization), the weed species diversity and richness and weed community evenness were the lowest. The principal component analysis showed that the weed community was divided into three groups: (i) NK and a part of NOF; (ii) NP and NPK; and (iii) PK and NOF.The correlation analysis indicated that the inuence of each macro-element on the weed community followed the model N > P > K. The organic content in the paddy soil might have played an equally important role with the amount of available N in determining the weed communitys characteristics. Regarding the way by which N, P, and K inuenced the weed community, the amount of available P and K mainly inuenced the organic content, while the amount of available N inuenced both the organic content and light transmittance within the canopy, thereby enhancing the capacity of rice to compete with weeds. Keywords: long-term fertilization, macro-elements, paddy eld, rice, weed biodiversity.

INTRODUCTION Recent emphasis has been placed on enhancing agricultural sustainability through a greater reliance on the ecological environment of the eld. In this context, the maintenance of biodiversity is viewed as an important coping strategy or an insurance against agricultural risks in an uncertain future (Loreau et al. 2003; Jackson et al. 2007). Arable weeds are important components of agroecosystems (Marshall et al. 2003) and play a vital role in supporting biological diversity (Hawes et al. 2003; Marshall et al. 2003; Gibbons et al. 2006).They constitute the base of the food chain for herbivores and their natural

Communicated by G.Wang. *Correspondence to: Fang Chen, Key Laboratory of Aquatic Botany and Watershed Ecology,Wuhan Botanical Garden, Chinese Academy of Sciences,Wuhan, 430074, China. Email: fchen@ipni.ac.cn The authors have no commercial interest in the ndings that are presented. This author contributed equally with the rst author to the article. Received 24 March 2011; accepted 10 December 2011

Published 2012. doi:10.1111/j.1445-6664.2011.00430.x This article is a U.S. Government work and is in the public domain in the USA.

Fertilization and weed biodiversity enemies, supporting many species of benecial insects, especially crop pollinators (Rypstra et al. 1999). Meanwhile, a high weed diversity is conducive to maintaining and regulating the microbial diversity of the soil and reducing the effects of harmful weeds. Some studies have suggested that there might be tighter N cycling and less nitrate leaching with increased weed diversity (Tilman et al. 1996; Scherer-Lorenzen et al. 2003). In a word, to maintain weed diversity in farmland is favorable to improving the yield and quality of crops and can ensure the sustainable utility of agro-ecosystems. However, in recent decades, because the transition from traditionally extensive to intensive agriculture changed the agricultural ecosystem, the number of arable weeds in farmland decreased sharply.This led to many intractable problems in relation to agricultural sustainability. The species richness pattern of the farmland weed community is very complex and is inuenced by many interactive factors, but ultimately it is attributed to specically agricultural practices (e.g. tillage, rotation, fertilization, and weeding; Froud-Williams 1988) to a large degree. Fertilization is an important agricultural measure that can improve the fertility level of the soil and result in new selective pressures to weed species and thus change the occurrence frequencies and community components of the weed seed bank (ODonovan et al. 2007; Andreasen & Skovgaard 2009; Smith et al. 2010). While improving the crops yield and quality, fertilization profoundly inuences the diversity of the whole weed community and its individual components (Dvorzak 1994; Theaker et al. 1995; Moss et al. 2004). Rice is one of the most widely cultivated crops in the world; thus, rice paddies are the main component of many agricultural landscapes. Therefore, the scientic management of paddy eld weeds has important signicance for the maintenance of healthy agro-ecosystems and the sustainable development of rice production. At present, the research about weed management in paddy elds mainly has converged on weed control, most of which has been by herbicides (Singh et al. 2006; Ishaya et al. 2007; Norsworthy et al. 2010), followed by biochemicals and allelochemicals (Ahn et al. 2005; Yu et al. 2005; Batish et al. 2007).The ideology of the research still has been in terms of regarding weeds as problematic living things that could be resolved by curative tactics. Actually, these curative tactics, especially the application of herbicides, already have brought forth many ecological and health problems for a long time.Therefore, in the past decade or more, researchers have started to regard weed management as a component of integrated cropping system design and have attempted to develop integrated weed management strategies (Mortensen et al.

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2000). The integrated weed management strategy needs to be underpinned by a signicant amount of knowledge of weed ecology and biology in agro-ecosystems (Wilson et al. 2009) and needs to incorporate multiple tactics, such as the prevention, avoidance, monitoring, and suppression of weeds, rather than simply reacting to weed infestations (Buhler 2002; Baumgartner et al. 2008). Nevertheless, developing integrated weed management systems that address a diversity of weed species with a diversity of life history traits is difcult.As a result of this diversity, a robust system requires ecological insight into the weed community and an understanding of how tillage practices complement one another in order to maintain weed populations at low equilibrium densities and to reduce the relative tness of the weeds: in other words, not only to prevent infestations of harmful weed species, but also to preserve weed biodiversity and thus maintain a healthy agro-ecosystem. Nowadays, although many reports about integrated weed management have been seen (Chikoye et al. 2004; Norsworthy & Frederick 2005; Chikowo et al. 2009), those about rice paddy elds are less numerous. This study investigated the impacts of long-term different types of fertilization on the weed biodiversity in rice paddy elds with red soil.The objective of this study was to comprehend how the weed community in the rice paddy eld would respond to different fertilization treatments and nutrition elements and ultimately to provide a scientic basis for developing an integrated weed management strategy in rice paddy soil. MATERIALS AND METHODS Experimental site and treatments The study was conducted in Long-term-located Monitoring Station for Soil Fertility, Agricultural Science Academy, of Jiangxi Province, China, at which a longterm fertilization experiment has lasted for 26 years.The crop rotation system was early rice late rice. Here, the annual temperature is 17.5C, the annual precipitation rate is 1600 mm, and the mean annual evaporation rate is 1800 mm.The cumulative temperature (rising at 10C increments) is 5400C and the number of frost-free days is 280. Cement ridges with a width of 50 cm and a depth of 45 cm were made to separate the experimental plots. They formed the independent irrigation channel of each plot. Five experimental treatments were set: (i) no fertilization (NOF); (ii), PK; (iii) NP; (iv) NK; and (v) NPK. The plot size was 20 m2, with three replications and randomly arranged.The application rates of N, P, and K were: N (urea) at 180 kg ha-2, P2O5 at 60 kg ha-2, and

Published 2012. This article is a U.S. Government work and is in the public domain in the USA.

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K.Wan et al. of shared species between Community A and Community B). The importance values of each species in each plot were calculated as per the formula: importance value = 1/3(relative density + relative frequency + relative coverage). The importance values of 11 common species in 15 plots were used as the original matrix for the principal component analysis (PCA) and cluster analysis (Fig. 1). The 15 experimental plots were plotted in an ordination diagram in which the bidimensional space was represented by the rst two canonical variables (i.e. Factor 1 and Factor 2) of the PCA (Fig. 2). The correlations between N, P, and K and factors 1 and 2, between N, P, and K and the biodiversity indices, and between light transmittance and the biodiversity indices and rice

K2O at 150 kg ha-2, respectively. These recommended fertilization rates were based on local farmers common practice and soil test results. The available P in the soil was not decient, as was the available N and K in the soil in this experimental eld, so less P fertilizer was used, compared with the N and K fertilizers.Whole P2O5 and 50% urea were used as the base fertilizer, while 25% urea was used for the top dressing at the tillering stage and another 25% urea was used for the heading stage. Also, 50% K2O was used at the tillering stage and another 50% K2O was used for the heading stage. Weed survey and data processing The weed survey was conducted in late September 2009 (the weeds owering and fruit initiation period and rices grain-lling period). A ve-point sampling method was used in the survey: ve quadrats each with a 0.25 m2 area being set in each plot. The weed species and number in the quadrats were recorded.The amount of light transmittance within the canopy was calculated from 10 measures of light intensity per plot, while each of the measures was the average of two numerical readings that were derived from the soil surface and the crop canopy. The rice was harvested and its yield was measured in early November 2009. Soil samples were taken in each plot to analyze the amount of available N, P, and K in the soil. The biodiversity of the weeds was measured, including: the species richness S (i.e. the amount of species included in a quadrat); the species diversity, which was measured by the Shannon-Wiener index (i.e. H = SPi lnPi, in which Pi is the proportion of individual numbers of the i species to the total individual number of each species in the quadrat. It was calculated from the formula as Pi = Ni/N, of which N is the total individual number of each weed species and Ni is the individual number of the i species); the degree of community dominance, as measured by the Simpson index, D = SPi2; the community evenness, as measured by the evenness index, J = H /lnS; the difference in the community composition, which was measured by the Whittaker index, bw = S/ma 1, in which ma is the average number of species; and the community similarity, as measured by the Srensen index, which is a qualitative analysis according to Cs = 2j/(a + b), and as measured by the BrayCurtis index, a quantitative analysis according to CN = 2jN/(aN + bN) ( j is the amount of shared species between Community A and Community B; a and b are the total amount of species of Community A and Community B, respectively; aN and bN are the total individual numbers of Community A and Community B, respectively; and jN is the minimal sum of individual numbers

Fig. 1. Cluster analysis of different treatments, based on the BrayCurtis index. NOF, no fertilization.

Fig. 2. Scatter diagram of 15 plots on the rst two canonical variables of the principal component analysis. ( ), No fertilization; ( ), PK; ( ), NP; ( ), NK; ( ), NPK.

Published 2012. This article is a U.S. Government work and is in the public domain in the USA.

Fertilization and weed biodiversity biomass also were tted.The data analysis was conducted with SPSS 16.0 (SPSS: An IBM Company, Chicago, IL, USA). RESULTS Soil nutrition, light transmittance, and rice biomass Table 1 displays the differences in the soil nutrients, light transmittance, and rice biomass in the various treatments. The amount of available N in the soil in the NK and zero-N fertilization treatments was signicantly lower than in the other treatments (P 0.05).The amount of available P in the soil in the zero-P fertilization treatment was signicantly lower than in the other treatments (P 0.05) and the amount of available K in the soil in the zero-K fertilization treatment was signicantly lower than in the other treatments (P 0.05).The N, P, and K content in the NOF treatment was relatively low. The Weed community components and structure

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amount of light transmittance decreased, according to the model NOF > PK > NP > NK > NPK. The rice biomass increased, as per the model NOF < PK < NK < NP < NPK.

Table 2 displays the differences in the weed species that occurred in the various treatments. A total of 11 species was discovered in this study, of which six species (i.e. Rotala indica, Monochoria vaginalis, Echinochloa crus-galli, Paspalum distichum, Cyperus difformis, and Lindernia procumbens) occurred in all ve treatments, while ve species (i.e. Marsilea quadrifolia, Eleocharis dulcis, Ludwigia prostrata, Tradescantia albiora, and Eleocharis yokoscensis) only occurred in some of the treatments. The weed community components in the different treatments were various. The dominant species in the NOF treatment (importance value of 1.5) were

Table 1. Differences in soil nutrition, light transmittance, and rice biomass between the treatments Characteristic Organic matter (g kg-1) 20.2 88.7 Available N (mg kg-1) Available P (mg kg-1) 5.9 32.3 Available K (mg kg-1) Light transmittance (%) 19.3 Rice biomass (kg ha-1) 8475 NOF 0.35c 22.8 0.67c 86.9 0.7d 65.6 1.12d 82.2 0.79a 19.1 238.50d 9840 PK 0.31b 25.7 0.25d 133.6 0.75a 61.9 0.82a 26.1 0.42a 10.5 334.50c 10 620 NP NK 0.55a 22.6 0.55a 83.2 0.80b 3.9 0.42e 74.4 0.62b 5.7 406.5b 10 110 NPK 0.70b 26.0 1.35e 127.4 0.45e 51.9 1.31b 62 0.19c 4.9 240bc 11 790 0.35a 0.91b 0.31c 0.75c 0.36c 443.50a

Same letters in the same row mean no signicant difference (P > 0.05), the different letters mean a signicant difference (P < 0.05), and the alphabetical order accords with the size order of the mean values (i.e. from large to small). NOF, no fertilization.

Table 2. Differences in the importance values of each species between the treatments Species Tradescantia albiora Ludwigia prostrate Eleocharis yokoscensis Marsilea quadrifolia Eleocharis dulcis Rotala indica Monochoria vaginalis Echinochloa crus-galli Paspalum distichum Cyperus difformis Lindernia procumbens 0.02 0.03 0.09 0.03 0.22 0.20 0.15 0.01 0.06 0.17 0.01 NOF 0.02b 0.02ab 0.03b 0.01ab 0.07b 0.02a 0.02a 0.00a 0.03b 0.02b 0.01a 0.01 0.04 0.15 0.05 0.08 0.21 0.14 0.04 0.11 0.11 0.06 PK 0.01b 0.02a 0.01a 0.02a 0.03c 0.01a 0.02a 0.02a 0.01b 0.02c 0.03a NP 0.00 0.00 0.00 0.02ab 0.03c 0.04c 0.06a 0.03a 0.06a 0.04d 0.01a 0.06 0.03 0.0 0.31 0.11 0.12 0.02 0.04 0.20 0.01 NK 0.03a 0.00 0.00c 0.04ab 0.02a 0.03b 0.01a 0.02a 0.02b 0.031a 0.02a 0.01 NPK 0.00 0.02ab 0.00 0.00 0.00 0.01d 0.04a 0.03a 0.09a 0.03d 0.06a

0.01 0.07 0.08 0.12 0.05 0.62 0.04 0.01

0.03 0.14 0.06 0.70 0.02 0.04

Same letters in the same row mean no signicant difference (P > 0.05), the different letters mean a signicant difference (P < 0.05), and the alphabetical order accords with the size order of the mean values (i.e. from large to small). NOF, no fertilization.

Published 2012. This article is a U.S. Government work and is in the public domain in the USA.

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K.Wan et al. available N (P < 0.05) and organic matter (P < 0.05), but a positive correlation with the amount of light transmittance (P < 0.05). Inuences of different types of fertilization on the biodiversity of the weed community Table 4 displays the differences in the biodiversity indices of the weed community under different types of fertilization. The underlined treatments in the following models imply no signicant difference between each other: species richness (PK NOF > NK > NP NPK); species diversity (PK NOF > NK > NP > NPK); community dominance (NPK NP > NK NOF PK); and community evenness (PK NOF NK> NP NPK). The species richness and diversity were high in the zero-N fertilization (PK and NOF) treatments. In the zero-P fertilization (NK) treatment, the species richness ranked at the middle level and the species diversity was higher. In comparison, under the zero-K fertilization (NP) and balanced fertilization (NPK) treatments, the species richness and species diversity were at their lowest. The level of community dominance was at its lowest, but the level of community evenness was high, in the PK treatment. In the NOF and NK treatments, the community dominance was low, while the community evenness

E. dulcis, R. indica, C. difformis, and M. vaginalis, while in the PK treatment, they were R. indica and E. yokoscensis. In the NP treatment, the dominant specie was P disti. chum, while in the NK treatment, the dominant species were C. difformis and E. dulcis. Finally, in the NPK treatment, the dominant specie was P. distichum. Therefore, the dominant species were P. distichum in the condition of a rich soil supply of N, R. indica in the condition of a poor soil supply of N, E. dulcis and C. difformis in the condition of a poor soil supply of P, and E. yokoscensis in the condition of a rich soil supply of P and K, while the importance values of M. vaginalis in all the treatments were larger. The scatter diagram (Fig. 2) indicated that, because of different types of fertilization, the weed community was divided into three groups: (i) NK and a part of NOF (low N; Table 1), mainly distributed in the positive direction of Factor 1; (ii) NP and NPK (high N; Table 1), mainly distributed in the negative direction of factors 1 and 2; and (iii) PK (high P and K and low N; Table 1) and NOF (low N, P, and K, Table 1), mainly distributed in the positive direction of Factor 2. Points (i) and (iii) overlapped with the NOF. Table 3 indicated that Factor 1 signicantly and negatively correlated with the amount of available N, available P, and organic matter (P < 0.05), while Factor 2 had signicant negative correlations with the amount of

Table 3. Detectable signicant correlations between the principal component analysis factors and soil nutrition and light transmittance Fitted curves Organic matter in the soil (g kg-1) Available N in the soil (mg kg-1) Available P in the soil (mg kg-1) Available K in the soil (mg kg-1) Light transmittance (%) Y = 3.4X + 11.9 (R2 = 0.27, P < 0.05)

Factor 1 Y = -1.5X + 23.5 (R2 = 0.40, P < 0.05) Factor 2 Y = -1.5X + 23.5 (R2 = 0.41, P < 0.05)
, no signicant correlation.

Y = -15.8X + 104.0 Y = -13.8X + 37.8 (R2 = 0.49, P < 0.01) (R2 = 0.24, P < 0.05) Y = -14.3X + 104.0 (R2 = 0.40, P < 0.05)

Table 4. Differences in the biodiversity indices of the weed community under various types of fertilization Index Species richness ShannonWiener index Simpson index Pielou index NOF 10.00 2.00 0.16 0.86 1.00ab 0.10ab 0.20b 0.03a 10.67 2.17 0.13 0.92 PK 0.58a 0.08a 0.01b 0.03a 6.33 1.25 0.42 0.68 NP 0.58c 0.13c 0.06a 0.05b 8.67 1.78 0.21 0.82 NK 0.58b 0.06b 0.02b 0.02a NPK 5.00 0.97 0.52 0.60 1.00c 0.23d 0.11a 0.08b

Same letters in the same row mean no signicant difference (P > 0.05), the different letters mean a signicant difference (P < 0.05), and the alphabetical order accords with the size order of the mean values (i.e. from large to small). NOF, no fertilization.

Published 2012. This article is a U.S. Government work and is in the public domain in the USA.

Fertilization and weed biodiversity was high. In contrast, under the NP and NPK treatments, the community dominance was at its highest, while the community evenness was at its lowest. The tted correlation relationships (see Table 5) indicated that the species richness, species diversity, and community evenness all were negatively signicantly correlated with the amount of available N in the soil (P 0.05), which had a signicant upward parabola that correlated with the amount of available P in the soil (P 0.05), signicantly negatively correlated with the amount of organic matter (P 0.05), and signicantly positively correlated with the amount of light transmittance (P 0.05).The amount of available K in the soil had signicant upward parabola correlations with species diversity and community evenness (P 0.05).Additionally, the community dominance was positively signicantly correlated with the amount of available N (P 0.05) and organic matter (P 0.05), whereas it negatively signicantly correlated with light transmittance (P 0.05).The organic content had a signicantly positive correlation with the amount of available N (P 0.05) and signicantly downward parabola correlations with the amount of available P and K (P 0.05; Fig. 3). Figure 4 shows that the light transmittance negatively correlated with the rice biomass and had a downward parabola that correlated with the amount of available N, but there was no signicant correlation with the amount of available P and K. Community similarity among the different treatments Both the values of the Srensen and BrayCurtis indices (Table 6) and of the cluster analysis (Fig. 1) consistently indicated that the weed community in the NOF, PK, and NK treatments, which were N-poor, were the most similar with each other, while the BrayCurtis index and cluster analysis also displayed that the weed community in the NPK and NP treatments, which were N-rich, were similar with each other. But, as a whole, the community similarities of the NPK treatment with the others were relatively low. The Whittaker index disclosed the following model: NPK (1.20) > NP (0.74) > NK (0.27) > NOF (0.10) > PK (0.03).This indicated that the order of species richness should be: NPK < NP < NK < NOF < PK. DISCUSSION AND CONCLUSION The results showed that fertilization had profound inuences on the weed community in the paddy eld. The soil nutrition variance resulted in large differences in the
Y = 0.0002X 2 - 0.02X + 1.30 (R2 = 0.40, P < 0.05)

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Table 5. Detectable signicant correlations between the light transmittance, nutrient and biodiversity indices of the weed community

Available N in the soil (mg kg-1) Organic matter in the soil (g kg-1) Light transmittance (%)

Y = 0.06X + 1.0 (R2 = 0.55, P < 0.01) Y = -0.02X + 0.5 (R2 = 0.46, P < 0.01) Y = 0.01X + 0.6 (R2 = 0.50, P < 0.01) Y = 0.27X + 4.9 (R2 = 0.57, P < 0.01) ShannonWiener index Simpson index Species richness Pielou index

Y = -0.20X + 5.6 (R2 = 0.65, P < 0.01) Y = 0.06X - 1.1 (R2 = 0.66, P < 0.01) Y = -0.04X + 1.8 (R2 = 0.62, P < 0.01) Y = -0.82X + 27.4 (R2 = 0.70, P < 0.01)
, no signicant correlation.

Published 2012. This article is a U.S. Government work and is in the public domain in the USA.

Fitted curves

Y = -0.02X + 3.6 (R2 = 0.77, P < 0.01) Y = 0.01X - 0.4 (R2 = 0.78, P < 0.01) Y = -0.05X + 1.3 (R2 = 0.76, P < 0.01) Y = -0.09X + 17.2 (R2 = 0.72, P < 0.01)

Y = 0.002X 2 - 0.10X + 2.43 (R2 = 0.70, P < 0.01) Y = -0.001X 2 + 0.04X + 0.02 (R2 = 0.74, P < 0.01) Y = 0.0004X 2 - 0.03X + 1.00 (R2 = 0.70, P < 0.01) Y = 0.007X 2 - 0.50X + 11.60 (R2 = 0.70, P < 0.01)

Available P in the soil (mg kg-1)

Y = 0.001X 2 - 0.1X + 3.50 (R2 = 0.40, P < 0.05)

Available K in the soil (mg kg-1)

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K.Wan et al.

Fig. 3. Detectable signicant correlations between the organic matter and the soil nutrition. ( ), available N (mg kg1); ( ), available P (mg kg1); ( ), available K (mg kg1).

Fig. 4. Detectable signicant correlations between the light transmittance and the soil nutrition and rice biomass. ( ), available N (mg kg1); ( ), rice biomass (RB) (kg ha1).

Table 6. Similarity indices in the weed community under long-term different types of fertilization NOF NOF PK NP NK NPK 1.00 0.84 0.95 0.78 PK 0.76 0.84 0.95 0.78 NP 0.41 0.48 0.89 0.80 NK 0.85 0.55 0.39 0.71 NPK 0.29 0.39 0.85 0.24 Whittaker index 0.10 0.03 0.74 0.27 1.20

Values above the diagonal that is formed by relate to the BrayCurtis index, while those below relate to the Srensen index.

Published 2012. This article is a U.S. Government work and is in the public domain in the USA.

Fertilization and weed biodiversity species diversity of the weed community.The degree of inuence of the different nutrition elements on the weed community varied. Amount of available nitrogen The following aspects were disclosed. First, the divergence results of the weed community under the different fertilization treatments showed that the low-N plots (NK and NOF) mainly distributed in the positive direction of Factor 1, whereas those with high N (NP and NPK) mainly distributed in the negative direction of factors 1 and 2 (Fig. 2). Second, in the case of the NK, PK, and NOF treatments, the species richness and diversity were at their highest and thus the community evenness was high; whereas, the community dominance was at its lowest. However, in the case of the NP and NPK treatments, the species richness and diversity were low, which brought forth the high community dominance, but low community evenness.Third, the amount of available N in the soil had negatively signicant correlations with the species richness and diversity and community evenness (P 0.05) and a positively signicant correlation with the community dominance (P 0.05; Table 5). Fourth, the species richness order that was disclosed by the Whittaker index (i.e. PK NOF > NK > NP NPK) indicated that the level of species richness in both the NP and NPK treatments (with high N availability in the soil) was low (Table 6). Fifth, the above four points were conrmed by a cluster analysis (Fig. 1) and the Srensen index (Table 6).All these were sufcient to conclude that a high level of available N in the soil favored the healthy development of the weed community, unlike a low level of available N in the soil. Nitrogen fertilization in the paddy eld could inuence not only the species richness and diversity, but also the weed population size, and consequently it could change the level of community evenness. An increase in the amount of available N in the soil could promote the occurrence of some dominant species and could lead to the decrease of species richness and diversity and the increase of species simplication. Some previous studies have shown similar results. For instance, Tilman (1987) reported that 60% of the species were replaced in high-N plots during secondary succession in abandoned elds. Pyek and Lep (1991) also revealed that both the dose and type of N fertilizer have a signicant effect on the composition of the weed community.The species richness decreased signicantly with an increased dosage of N. Second, in light of the correlations between the available N, P, and K in the soil and factors 1 and 2, Factor 1 had a negatively signicant correlation with the amount

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of available N (P = 0.0037) and P (P = 0.0491) (See Table 3), while Factor 2 had a negatively signicant correlation with only the amount of available N (P < 0.05). No signicant correlation between the amount of available K and both factors 1 and 2 was detected. This suggests that the level of inuence of a single element on the weed community structure was exhibited as per the model N > P > K. Amount of available phosphorus and potassium The amount of available P and K in the soil had signicant upward parable correlations with the species richness, species diversity, and community evenness and a signicant downward parable correlation with the community dominance (P 0.05, Table 5), which suggests that, similarly to N, these two elements could inuence not only the species richness and diversity, but also the weed population size, and consequently they could change the community evenness. The correlations between the organic content in the soil and the parameters of species richness, species diversity, community evenness, community dominance, and factors 1 and 2 from the PCA were entirely consistent with those between the amount of available N in the soil and the parameters (Tables 3,5), suggesting that the organic content in the paddy soil plays an equally important role with the amount of available N in the soil in determining the weed communitys characteristics. The species richness, species diversity, and community evenness all signicantly correlated with the amount of available N, P, and K, organic matter content, and light transmittance (Table 5). However, because the organic content had a signicant correlation with the amount of available N, P, and K (Fig. 3), whereas the light transmittance only had a signicant correlation with the amount of available N (Fig. 4), therefore in relation to the way by which N, P, and K regulated the weed community, the amount of available P and K mainly inuenced the organic content, while the amount of available N inuenced both the organic content and the light transmittance (derived from inuencing the rice biomass), thereby enhancing the capacity of rice to compete with weeds. In relation to these results, it is puzzling that the amount of available K in the soil signicantly correlated with species diversity, species richness, and community evenness (Table 5), but nevertheless its signicant correlations with factors 1 and 2 were not detected (Table 3). This needs further exploration. According to the above analysis, a reasonable explanation can be made regarding the weed communitys characteristics in Table 4 under different types of fertilization,

Published 2012. This article is a U.S. Government work and is in the public domain in the USA.

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K.Wan et al. REFERENCES

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combined with the soil nutrient status in Table 1. In the PK treatment, which possessed a low N content and a high P and K content, due to the absence of N, the rice biomass was constrained (Fig. 4). Therefore, the light transmittance improved. Moreover, a high P and K content also elevated the weed biodiversity level, so that the species richness, species biodiversity, and community evenness were high, while the community dominance was low.The Simpson index also indicated that dominant species did not occur in the weed community. Likewise, in the NK and NOF treatments, although the K and P content in the soil exhibited different traits, the amount of available N in the soil was similarly low.The suppressed rice growth provided relatively benecial habitats for the weed species.Therefore, the species richness and diversity were also high, while the community evenness was high and thereby the community dominance was low. In the NP and NPK treatments, although the P and K content in the soil displayed different characteristics, the amount of available N was high. The high N content promoted the growth of rice and tall nitrophilous weeds. As a result of canopy shading, the light transmittance was largely weakened. Therefore, the light-loving species were strongly restrained and thus the community mainly consisted of fewer tall species and shade-tolerant species. In both these treatments, the species richness, species diversity, and community evenness were low, while the species dominance was high. The Simpson index also suggested that dominant species occurred in these communities. It should be pointed out that conceptions of integrated weed management and contemporary agronomy suggest that the regulation of the main components of the agrophytocenosis should diminish the number of weed populations to both economically and ecologically acceptable levels (Augustin 1989; Hain 1997). Nevertheless, the results in this study apparently showed that, in the case of balanced fertilization, the rice yield was assured, but that the level of weed biodiversity was very low.According to these results and in view of the importance of crop edges in preserving the weed biodiversity (Romero et al. 2008), the authors suggest that it is very necessary in the future to further study the role that is played by paddy eld edges in weed biodiversity conservation. ACKNOWLEDGMENTS This research project was funded by the Program of International Plant Nutrition Institute (IPNI-HB-33, 34), Wuhan, China and the Opening Project of Hubei Key Laboratory of Wetland Evolution & Ecological Restoration.

Published 2012. This article is a U.S. Government work and is in the public domain in the USA.

Fertilization and weed biodiversity

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Published 2012. This article is a U.S. Government work and is in the public domain in the USA.