ALLELOPATHIC EFFECT of INVASIVE SPECIES GIANT GOLDENROD (SOLIDAGO GIGANTEA AIT.) on CROPS and WEEDS* Renata Baličević, Marija

Herbologia, Vol. 15, No. 1, 2015

DOI 10.5644/Herb.15.1.03

ALLELOPATHIC EFFECT OF INVASIVE SPECIES GIANT GOLDENROD (SOLIDAGO GIGANTEA AIT.) ON CROPS AND WEEDS*

Renata Baličević, Marija Ravlić, Tea Živković* Faculty of Agriculture, Josip Juraj Strossmayer University of Osijek, Kralja Petra Svačića 1d, 31000 Osijek, Croatia, corresponding author: [email protected] *Student, Graduate study

Abstract The aim of the study was to determine allelopathic potential of invasive species giant goldenrod (Solidago gigantea Ait.) on germination and initial growth crops (carrot, barley, coriander) and weed species velvetleaf (Abutilon theophrasti Med.) and redroot pigweed (Amaranthus retroflexus L.). Experiments were conducted under laboratory conditions to determine effect of water extracts in petri dish bioassay and in pots with soil. Water extracts from dry aboveground biomass of S. gigantea in concentrations of 1, 5 and 10% were investigated. In petri dish bioassay, all extract concentrations showed allelopathic effect on germination and seedling growth of crops with reduction over 25 and 60%, respectively. Both weed species germination and growth were greatly suppressed with extract application. In pot experiment, allelopathic effect was less pronounced. Reduction in emergence percent, shoot length and fresh weight of carrot were observed. Barley root length and fresh weight were reduced with the highest extract concentration. No significant effect on seedling emergence and growth of A. theophrasti was recorded, while emergence of A. retroflexuswas inhibited for 14.4%. Germination and growth of test species decreased propor- tionately as concentration of weed biomass in water extracts increased. Differences in sensitivity among species were recorded, with A. retroflexus being the most susceptible to extracts. Key words: allelopathy, water extracts, Solidago gigantea, germination, crops, weeds

Introduction Excessive application of herbicides in most weed management sys- tems is a major concern since it causes serious threats to the environment, public health and increases costs of crop production. Weed seed germina-

Copyright © 2015 by the Academy of Sciences and Arts of Bosnia and Herzegovina. Baličević et al. tion inhibition and growth suppression which can be attributed to allelopathy is highly important and can be considered as a possible alterna- tive, non-chemical weed management strategy (Asghari and Tewari, 2007, Macias, 1995). Allelopathy represents the influence of one organism on the other, whether it is harmful or beneficial, through production of allelochemicals (Rice, 1984). Allelopathic crops for controlling weeds could be used either directly as cover, smother or green manure crops or by using allelochemicals as natural herbicides (Singh et al., 2003). Currently, aromatic and medicinal plants are investigated as potential allelopathic crops (Đikić, 2005, Baličević et. al., 2014, Ravlić et al., 2014). Beside crops, weed species with high inhibitory effect also have the potential to be used in control of other weeds (Qasem and Foy, 2001, Galzina et al., 2011). Giant goldenrod (Solidago gigantea Ait.), species belonging to the family Asteraceae, is a perennial herb native to North America and in- troduced in Europe as an ornamental plant (Weber and Jakobs, 2005). S. gigantea is an invasive species which prefers ruderal habitats, river- sides, forests, roadsides etc. Its number and aggressiveness is extensive on abandoned agricultural areas where it eliminates indigenous plants, however it is rarely found on agricultural fields (Knežević, 2006, Novak and Kravarščan, 2011). Allelopathic potential of goldenrod extracts on different crops was reported previously (Sekutowski et al., 2012, Beres and Kazinczi, 2000). Bortniak et al. (2011) states that emission of allelopaths to soil by S. gigantea may be a potential threat in case of reinstating waste- land colonized by this species for agricultural production. Since harmful effect of allelopathic plant has to be targeted on the weeds, while at the same time the crops must be unaffected or tolerant, it is important to assess its potential on wide range of species, both crops and weeds. The aim of the study was to determine allelopathic effect of water extracts from dry biomass of S. gigantea on germination and growth of crops (carrot, barley, coriander) and weed species velvetleaf (Abutilon theophrasti Med.) and redroot pigweed (Amaranthus retroflexus L.) both in petri dishes and pots with soil.

Materials and methods Plants of giant goldenrod (S. gigantea) were collected in late sum- mer of 2014 at the flowering stage (Hesset al., 1997) from ruderal habitats (edges of farm fields). Aboveground fresh biomass was oven dried, and after that chopped into small pieces. Dried pieces were ground with elec- tronic grinder into fine powder.

20 Allelopathic effect of invasive species giant goldenrod (Solidago gigantea Ait.)

Water extract was prepared according to Norsworthy (2003) by mix- ing 100 grams of dry goldenrod biomass with 1000 ml of distilled water and kept for 24 h at room temperature. The mixture was filtered through muslin cloth to remove debris and after that through filter paper. The ob- tained extract was diluted with distilled water to give final concentrations of 1, 5 and 10% (10, 50 and 100 g/l). Winter wheat (cv. Barun) was acquired from Agricultural Institute Osijek, while carrot and coriander seeds were purchased from seed com- pany. Weed seeds of velvetleaf (A. theophrasti) and redroot pigweed (A. retroflexus) were collected during 2014 from agricultural fields in Osijek- Baranja County. All seeds used in experiments were surface-sterilized for 20 minutes with 1% NaOCl (4% NaOCl commercial bleach), then rinsed three times with distilled water (Siddiqui et al., 2009). Effect of goldenrod extract was determined in two set of experiments: i) petri dishes bioassay with filter paper and ii) pot culture with soil. In petri bioassay, effect of three concentrations of extract (1, 5 and 10%) was evaluated. Twenty five (barley, coriander) or thirty (carrot, velvetleaf, redroot pigweed) seeds were placed in sterilized Petri dishes (90 mm) lined with filter paper. In each petri dish an equal amount of certain extract was added (5 ml for barley, coriander, velvetleaf and 2 ml for carrot and redroot pigweed), while distilled water was used in control. Additional extract/water was added to prevent seeds from drying. Each treatment had four replications, and experiment was repeated twice. Petri dishes were kept at room temperature (22°C ± 2) for 8 days. In experiment with pots, effect of two concentration of extract (5 and 10%) was evaluated. Thirty seeds of crop or weed were sown in pots (9x7 cm for barley and velvetleaf; 9x4 cm for carrot and pigweed) filled with commercial substrate (NPK 210:120:260 mg/l, pH 5.6). Each pot was watered with 75 ml of extract per 100 g of soil, while distilled water was used in control. Thereafter, all treatments were equally watered. Redroot pigweed was grown for 7, barley and velvetleaf for 12, and carrot for 14 days under the same conditions on laboratory benches at 22°C ± 2 temperature. All treatments had four replications and all experiments were conducted twice. Germination percentage was calculated as G (Germination,%) = (Germinated seed/Total seed) x 100. All emerged seedlings were counted and percentage of emergence was calculated as E= (Emerged seed/Total seed) x 100. At the end of each experiment, seedling root length (cm), shoot length (cm) and fresh weight (g, mg) were determined. The collect-

21 Baličević et al. ed data were analysed statistically with ANOVA and differences between treatment means were compared using the LSD-test at probability level P<0.05.

Results and discussion S. gigantea water extract had various effect on germination of crops in petri dish bioassay (Figure 1). Germination was inhibited in all test species, however only higher concentrations showed significant effect. Carrot seed germination was reduced for up to 35.9%, and barley germination for up to 38.8%.

Figure 1. Effect of S. gigantea water extracts on germination of crops (petri dish bioassay)

Similarly, seedling growth of crops was differently affected with the extract application (Table 1 and Table 2). Coriander seedling length and fresh weight was significantly reduced with the highest extract concentration, while lower concentrations of extract stimulated coriander seedling growth.

22 Allelopathic effect of invasive species giant goldenrod (Solidago gigantea Ait.)

Table 1. Effect of S. gigantea water extracts on seedling growth of coriander (petri dish bioassay) S. gigantea Root length Shoot length Fresh weight biomass in (cm) (cm) (g) water, g/l Coriander 0 5.41 b 4.02 b 0.026 b 10 6.90 a 5.62 a 0.033 a 50 3.45 c 4.01 b 0.024 b 100 0.19 d 1.73 c 0.011 c Means followed by the same letter within the column are not significantly different at P<0.05.

Root and shoot length and fresh weight of carrot were significantly decreased only with higher extract concentrations. Seedling length and fresh weight of barley was greatly inhibited. The lowest concentration reduced root and shoot length for 36.4 and 22.1%, respectively, while reduction in treatment with the highest concentration amounted up to 75.2%.

Table 2. Effect of S. gigantea water extracts on seedling growth of carrot and barley (petri dish bioassay) S. gigantea Root length Shoot length Fresh weight biomass in (cm) (cm) (g) water, g/l Carrot 0 1.66 b 1.99 a 0.0041 a 10 2.08 a 2.05 a 0.0037 a 50 1.61 b 1.32 b 0.0022 b 100 0.36 c 0.35 c 0.0003 c Barley 0 11.03 a 9.34 a 0.215 a 10 7.02 b 7.28 b 0.136 b 50 3.48 c 3.59 c 0.065 c 100 2.74 c 2.55 d 0.049 c Means followed by the same letter within the column (for each crop) are not significantly different at P<0.05.

23 Baličević et al.

Germination of weed species A. theophrasti and A. retroflexus was significantly inhibited with extract in all concentrations and reduction was up to 71.8 and 97.4%, respectively (Figure 2).

Figure 2. Effect of S. gigantea water extracts on germination of weeds (petri dish bioassay)

Goldenrod water extract showed both negative and positive effect on growth of weeds in petri dish bioassay (Table 3). Reduction in root and shoot length and fresh weight of A. theophrasti seedlings was observed with all extract concentrations. The highest concentration inhibited above- mentioned parameters for 89.7, 83.2 and 81.4%, respectively. Both higher concentrations showed significant negative effect on seedling growth of A. retroflexus. Reduction of root length ranged from 97.4 to 98.3%, while shoot length decreased for over 91.9%. Significant decrease in fresh weight of weed seedlings was also recorded. Contrary, lower concentration of extract had significant stimulatory effect on seedlings growth. Shoot length of weed was greater for 53.8%, and fresh weight for 26.9%, compared to the control treatment.

24 Allelopathic effect of invasive species giant goldenrod (Solidago gigantea Ait.)

Table 3. Effect of S. gigantea water extracts on seedling growth of weeds (petri dish bioassay) S. gigantea Root length Shoot length Fresh weight biomass in (cm) (cm) (mg) water, g/l Abutilon theophrasti 0 4.17 a 4.69 a 68.63 a 10 3.52 b 3.94 a 54.38 b 50 1.03 b 1.53 b 29.10 c 100 0.43 b 0.79 b 12.75 d Amaranthus retroflexus 0 3.09 a 1.99 b 4.71 b 10 3.54 a 3.06 a 5.98 a 50 0.05 b 0.16 c 0.31 c 100 0.08 b 0.14 c 0.08 c Means followed by the same letter within the column (for each weed) are not significantly different at P<0.05.

Table 4. Effect of S. gigantea water extracts on emergence and seedling growth of carrot and barley (pot experiment) S. gigantea Emergence Root length Shoot length Fresh weight biomass in (%) (cm) (cm) (g) water, g/l Carrot 0 57.9 a 2.69 a 3.67 a 0.010 a 50 46.7 b 2.15 a 2.87 b 0.008 b 100 39.6 b 2.37 a 2.34 b 0.007 b Barley 0 99.0 a 12.28 a 19.91 a 0.393 b 50 98.0 a 12.56 a 20.48 a 0.433 a 100 98.0 a 10.72 b 20.14 a 0.365 c Means followed by the same letter within the column (for each crop) are not significantly different at P<0.05.

Application of extract to pots with soil resulted in emergence reduction of carrot seedlings for up to 31.6% (Table 4). Seedlings shoot length

25 Baličević et al. and fresh weight was similarly affected. Extracts had no significant effect on root length of carrot seedlings. Emergence of barley was not affected when goldenrod extracts were applied. Root length and fresh weight of barley was significantly reduced with the highest extract concentration, for 12.7 and 7.1%. Lower concentration of extract had positive effect on barley growth. Extract had no significant effect on seedling emergence and growth of A. theophrasti, although shoot length and fresh weight were suppressed with higher extract concentration (Table 5). Extracts stimulated seedling length and fresh weight of A. retroflexus, while emergence was inhibited with 10% concentration for 14.4%.

Table 5. Effect of S. gigantea water extracts on emergence and seedling growth of weeds (pot experiment) S. gigantea Emergence Root length Shoot length Fresh weight biomass in (%) (cm) (cm) (g) water, g/l Abutilon theophrasti 0 78.7 a 4.84 a 9.28 a 0.105 a 50 78.3 a 4.73 a 9.30 a 0.106 a 100 79.6 a 5.09 a 8.94 a 0.102 a Amaranthus retroflexus 0 65.8 ab 2.21 a 4.04 b 0.0073 b 50 67.1 a 2.57 a 5.02 a 0.0089 a 100 56.3 b 2.33 a 4.42 b 0.0076 b

Means followed by the same letter within the column (for each weed) are not significantly different at P<0.05.

Extracts from S. gigantea showed allelopathic potential, both stimulatory and inhibitory, in petri dishes and in pot experiments. This is in agreement with results of Sekutowski et al. (2012) who recorded reduced germination and root length of buckwheat, and stimulation of root growth of sunflower when goldenrod fresh biomass extracts were applied in petri dishes. Bortniak et al. (2011) also investigated effect of fresh extracts from roots and leaves of goldenrod under greenhouse conditions. The results indicated that inhibitory potential was greater against winter wheat and triticale, while oilseed rape, barley and rye germination and root length were less affected.

26 Allelopathic effect of invasive species giant goldenrod (Solidago gigantea Ait.)

The results showed extracts had greater effect on seedling growth of crops rather than on germination. This is in agreement with findings of others (Konstantinović et al., 2014; Baličević et al., 2014b) who also concluded that effect of allelochemicals can be more pronounced on the growth of seedlings. Crops and weeds differed in their susceptibility to S. gigantea water extracts, in petri dishes and in pots. In petri dishes, barley was the most susceptible species among crops with average seedling growth reduction over 50%. Both weed species were more sensitive to extracts applied, and A. retroflexus seed germination and growth were inhibited over 85 and 40%, respectively. In pots, carrot emergence and seedling growth were the most affected. Sensitivity of crops and weeds to allelochemicals is different among species and genotypes within species (Asghari and Tewari, 2007, Baličević et al., 2014b). Generally, germination and growth of all test species decreased pro- portionately as concentration of weed biomass in water extracts increased. In addition, lower concentration showed stimulatory effect on growth of certain species. Inhibitory effect of higher and stimulatory effect of lower concentrations of weed water extracts was also reported by Marinov- Serafimov (2010) and Baličević et al. (2014). However, differences between results from experiments in petri dish and pots were observed and allelopathic effect was altogether more pronounced in petri dish bioassay. Higher inhibitory effect of water extracts could be due to direct contact of seed with extract on filter paper or adsorption of allelochemicals to soil. Adsorption of allelochemicals to soil can reduce its effect, especially in agro-ecosystems (Vidal et al., 1998).

Conclusions The results of the experiments showed that extracts from dry goldenrod biomass had certain allelopathic effect on both crops and weeds. Further research, in laboratory and field, towards determination of different doses and application time of goldenrod extract, as well as finding tolerant crops and susceptible weeds is in need to fully assess goldenrod allelopathic potential.

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